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BULLETIN OF 

THE BRITISH MUSEUM 

(NATURAL HISTORY) 



ZOOLOGY 
VOL. VI 
1959-1960 




PRINTED BY ORDER OF THE TRUSTEES OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 
london: i960 



DATES OF PUBLICATION OF THE PARTS 



No. 


i. 


23 October 1959 


No. 


2. 


19 February i960 


No. 


3- 


11 March i960 


No. 


4- 


11 March i960 


No. 


5- 


11 March i960 


No. 


6. 


12 April i960 


No. 


7- 


22 July i960 



PRINTED IN 

GREAT BRITAIN 

AT THE 

BARTHOLOMEW PRESS 

DORKING 

BY 

ADLARD AND SON, LTD. 



CONTENTS 

ZOOLOGY VOLUME 6 

PAGE 

No. i. Taxonomy of the genus Columba. By D. Goodwin i 

No. 2. A revision of the Platyseiinae (Mesostigmata: Aceosejidae) based on 
material in the collections of the British Museum (Natural History). 
By G. Owen Evans and K. H. Hyatt 25 

No. 3. British slugs (Pulmonata: Testacellidae, Arionidae, Limacidae). By 

H. E. Quick (Pis. 1-2) 103 

No. 4. A revision of the Lake Victoria Haplochromis species (Pisces, Cichlidae) 

Part IV. By P. H. Greenwood 227 

No. 5. On a Stygiomysis from the West Indies, with a note on Spelaeogriphus 

(Crustacea, Peracarida). By Isabella Gordon (Pis. 3-4) 283 

No. 6. Contributions toward a classification of modern Isospondylous fishes. 

By William A. Gosline 325 

No. 7. The ecology and taxonomy of some Angola birds (Based on a collection 

made in 1957). By B. P. Hall (Pis. 5-6) 367 

Index to Volume 6 455 




TAXONOMY OF THE GENUS 

COLUMBA 




D. GOODWIN 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ZOOLOGY Vol. 6 No. i 

LONDON: 1959 



TAXONOMY OF THE GENUS COLUMBA 



BY 

D. GOODWIN 



Pp. 1-23 ; 5 Text-figures 




BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ZOOLOGY Vol. 6 No. 1 

LONDON: 1959 



THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical Series. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

This paper is Vol. 6, No. 1 of the Zoological series. 



Trustees of the British Museum, 1959 



PRINTED BY ORDER OF THE TRUSTEES OF 
THE BRITISH MUSEUM 

Issued October, 1959 Price Nine Shillings 



TAXONOMY OF THE GENUS COLUMBA 

By D. GOODWIN 

CONTENTS 

Page 

Introduction ........... i 

Colour and Colour-pattern ........ i 

Adaptive Radiation and Relationships ...... 2 

The Species-groups and their Component Species .... 4 

Notes on Species .......... 7 

Summary ............ 22 

References ........... 22 

SYNOPSIS 
Previous reviews of the genus Columba have grouped together species which lack display 
plumage on the neck or whose ornamentation is superficially similar. This is believed to have 
resulted in some false conclusions because such display plumage has been independently lost 
or become obsolescent in several species. When this is recognized and greater importance 
attached to basic colour-patterns and geographical distribution a truer picture of relationships 
within the genus can be arrived at. 

INTRODUCTION 

The genus Columba includes a large number of medium to large-sized pigeons, 
widely distributed throughout the world. Peters (1937), for example, lists fifty-two 
species and does not divide them into sub-genera. The rather inconveniently large 
size of the genus has often been criticized and the affinities of its members questioned. 
Ridgway (1916) separated the relatively few northern and central American species 
into six genera, some of them monotypic. The most important recent revision of the 
genus is that of Boetticher (1954), who divides it into several sub-genera and corrects 
some false conclusions of a previous and less comprehensive revision by Miculicz- 
Radecki (1949). 

When re-arranging the very extensive collection of specimens of Columba in 
the British Museum (Natural History) it was found impossible to agree with all 
of Boetticher's conclusions. The object of this paper, therefore, is a revision of 
the genus based on examination of the skins in the National Collection and correlated, 
as far as has been possible, with what is known of the ecology and behaviour of 
the species of which it is comprised. Emphasis is placed on discussion of those 
points where my conclusions differ from those of Boetticher. I am indebted to Dr. 
Dean Amadon of the American Museum of Natural History for the loan of specimens 
of Columba albinucha and C. pallidiceps, to Mr. J. D. Macdonald for help and con- 
structive criticism and to Dr. G. Bodenstein, Professor W. F. Hollander and Mr. 
R. W. Sims for information and discussion on various points. 

COLOUR AND COLOUR-PATTERN 

Within Columba plumage colour ranges from entirely black, with green and purple 
iridescence, to predominantly pale grey or purplish-chestnut. It is evident that, 

ZOOL. 6. I. 1 



TAXONOMY OF THE GENUS COLUMBA 



as in Ptilinopus (Cain, 1954) colour-pattern is usually a better guide to relationship 
than actual colour. One finds striking differences in the colour of the plumage and/ 
or soft parts of the head and neck (the focal area of both sexual attraction and aggres- 
sive attack in pigeons) between pairs of sympatric and very closely related species 
as C. squamosa and C. leucocephala or C. rupestris and C. leuconota (see Text-fig. 1). 





ji'i'i'i'i'i! 
1,11,1 



rz-Z-Z-rz Slate-grey or blue-grey 



KEY TO SHADING 



Dull purple or purplish-brown 



(descent green 



^=r- *xWSc X Iridescent purple or wine-red 






Black or greyish-black, glossed with 
green in pallidiceps 



White or silver-grey 



Display plumage with dark feather- 
edges giving a scale-like appearance 



Fig. 1. Diagrammatic sketches to show differences in head and neck colours between 
three pairs of sympatric and closely related species. From top left : pallidiceps and 
vitiensis, squamosa and leucocephala, rupestris and leuconota. 

Such differences almost certainly function as isolating mechanisms to inhibit inter- 
specific pairing. Signal markings on wings and tail and specific differences in voice 
and display-flight serve the same purpose at " longer range ". As with other groups 
of birds the greatest diversity of signal markings, display plumage and voice is 
found where there are many related and at least potentially sympatric species and, 
conversely, a tendency to lose such characters and acquire a dull uniformly coloured 
plumage in isolated island forms. Lack of signal markings or display plumage in 
Columba seems never to be a primitive character. Examination of the colour- 
pattern and other characters of the species concerned shows convincingly that their 
common lack of ornamentation is due to convergence and not to close affinity. 

Sexual dimorphism in Columba is relatively slight but in most species it is sufficient 
to enable skins or living birds to be correctly sexed on plumage colour alone. In 
those species which are predominantly grey males are a purer, bluer grey than 
females. Purple, chestnut or mauve-pink in males is often duller, absent, or replaced 
by dull brown in females. Iridescent display plumage (when present) is usually more 
brilliant in males. Where adults show such differences of plumage similar and homo- 
logous sexual differences are found in the juvenile plumage. 

ADAPTIVE RADIATION AND RELATIONSHIPS 

Adaptive radiation can be misleading at generic level, since once two forms have 
evolved marked differences of size and structure they are unlikely to be considered 



TAXONOMY OF THE GENUS COLUMBA 3 

con-generic. For example the many Australian bronze- wing pigeons are obviously 
very close relatives in spite of the taxonomically convenient differences in size 
and proportions which they have developed in the course of adaptation to different 
ecological niches. Within Columba, as here recognized, the greatest amount of 
adaptive radiation is found among the Palearctic and Ethiopian species. Similar, 
but less marked differences are shown among the American species, but here maculosa, 
which otherwise closely parallels the old-world livia, has not emancipated itself 
from the use of trees for resting and breeding. 

The question arises whether the New World representatives of Columba should 
be generically separated from the Old World forms. Cumley & Irwin (1944) concluded 
from their work on the antigenic inter-relationships of the blood cells of guinea, 
livia, palumbus, janthina, fasciata, maculosa, picazuro, cayennensis, flavirostris and 
leucocephala that each of the six American species was more closely related to the 
other five than to any of the four Eurasian species. Verheyen (1957) considers that 
skeletal differences warrant the separation of Old and New World forms of Columba, 
both of which groups he further divides into several genera, some of them monotypic. 
I am not competent to criticize Verheyen's opinions in so far as they are based on 
osteology, but his decisions placing guinea in the monotypic genus Dialiptila and 
generically separating janthina, vitiensis and norfolciensis are so contrary to the 
evidence from external taxonomic characters and behaviour-patterns that I hesitate 
to follow his conclusions without further evidence. 

If the New World species are to be generically separated this could be done only 
on osteological and biochemical characters. Possibly studies currently in progress in 
America will show that there are constant and significant behavioural differences 
also and it will be desirable to separate the American from the Old World species. 
I think, however, that at present it would be premature to do so. 

It seems to me best to retain the large genus Columba as used by Peters (1937) and 
Hellmayr & Conover (1942) but to give Oenoenas and Turturoena sub-generic 
rank within it. I also think that Nesoenas should be given only sub-generic rank 
within Columba. The remaining forms can be relegated to five species-groups. 
Cain (1954), in his revision of the pigeons in the genus Ptilinopus, used the category 
of the species-group for " an informal natural group of closely related species below 
the rank of a subgenus, the recognition of which involves no complication of the 
scientific names of the species contained in it." And that of sub-group for 
more closely related forms within the species-group. I have followed Cain in the 
use of these categories which are equally useful for defining relationships of the 
pigeons in Columba. The hierarchy of categories is therefore : 

Genus 
Sub-genus ' 

Species-group 
Sub-groups (of species group) 
Superspecies 
Species 
Subspecies 

ZOOL. 6, I. I§ 



4 TAXONOMY OF THE GENUS COLUMBA 

There are, naturally, some species whose characters show them to be inter- 
mediate between two sub-groups and it has been necessary to allot them somewhat 
arbitrarily to that with which they seem to have most in common or else to place them 
in a mono-specific sub-group. All such cases are discussed below. In the 
following list members of the same superspecies are bracketed together. Sub- 
species have not been listed but they are discussed under the species headings where 
they are relevant to specific problems. 



THE SPECIES-GROUPS AND THEIR COMPONENT SPECIES 

(i) Oenas Species-group 

Smallish (oliviae) to large (trocaz) sized columbids inhabiting the Palearctic and 
Ethiopian regions, one species (livia) also occurring in India, and throughout much 
of the world in a feral state as a result of human introduction. Most species in this 
group have well developed display plumage on the neck and patterned tails. Sexual 
dimorphism slight. 

(a) Palumbus Sub-group 

Tail rather long with conspicuous pale central band ; breast mauve-pink or 
salmon-pink. 

C. palumbus Linnaeus, 1758 

C. trocaz Heineken, 1829 

C. bollii Godman, 1872 

C. unicincta Cassin, 1859 (i860) 

(b) Oenas Sub-group 

Glossless mauve-pink on head and /or breast, black signal markings on inner 
wing-coverts (obsolescent in oliviae). 



{ 



C. oenas Linnaeus, 1758 

C. eversmanni Bonaparte, 1856 

C. oliviae Stephenson Clarke, 191 8 



(c) Livia Sub-group 

No glossless mauve-pink colour on breast or head. Display plumage on front as 
well as sides of neck (absent in leuconota) . 



{ 



C. livia Gmelin, 1789 
C. rupestris Pallas, 181 1 
C. leuconota Vigors, 1831 
C. guinea Linnaeus, 1758 
C. albitorques Ruppell 1837 



TAXONOMY OF THE GENUS COLUMBA 5 

(2) Junioniae Species-group 

Large columbid inhabiting the western Canary Islands of Palma and Gomera. 

Mantle and wings dark brown and immaculate ; tail with broad pale terminal 
band. Entire neck and breast iridescent but no well differentiated display plumage. 
One species C. junioniae Hartert, 1916. 

(3) Elphinstonii Species-group 

Smallish medium to large-sized columbids inhabiting the Indo-malayan and Ethi- 
opian regions. All except one with unpatterned tails. No signal markings on wings. 
Most with display plumage on sides and back of neck. 

(a) Arquatrix Sub-group 

Display plumage on hind neck lanceolate. Wing-coverts and /or underparts 
spotted with white (obsolescent spots in polleni). 

CC. arquatrix Temmink, 1809 

< C. pollenii Schlegel, 1866 
\_C. hodgsonii Vigors, 1832 

C. albinucha Sassi, 1911 

(b) Elphinstonii Sub-group 

Display plumage on hind neck of black feathers tipped white or buff. Iridescent 
green or purple sheen on neck and mantle. Unpatterned tails, no signal markings 
on wings. One superspecies. 

CC. pulchricollis Blyth, 1845 (1846) 

< C. elphinstonii (Sykes, 1832) 
I^C. torringtoni Bonaparte, 1854 

(c) Punicea Sub-group 

No well differentiated display plumage on neck ; wing coverts and body plumage 
immaculate chestnut or silver contrasting with black primaries. 

C. punicea Blyth, 1842 

C. argentina Bonaparte, 1855 

(d) Palumboides Sub-group 

Head and neck silver-grey ; wings and back dark grey with iridescent feather 
edges. One species. 

C. palumboides (Hume, 1873) 

(4) Janthina Species-group 

Large columbids inhabiting the Pacific regions from Japan to Australia, chiefly 
on islands. Plumage largely or entirely blackish and highly iridescent. No signal 
markings on wings ; dark unpatterned tails. Not divisible into sub-groups. 



6 TAXONOMY OF THE GENUS COLUMBA 

CC. janthina Temminck, 1830 
< C. vitiensis Quoy & Gaimard, 1830 
[_C. norfolciensis Latham, 1801 

C. versicolor Kittlitz 

C. jouyi Stejneger, 1887 

C. pallidiceps (Ramsay, 1877) 

(5) Picazuro Species-group 

Small (rufina) to medium-large (maculosa) columbids inhabiting the nearctic and 
nootropic regions. Most with display plumage on back and sides of neck and patterned 
tails. 

(a) Leucocephala Sub-group 

Patch of velvet-like feathers on nape ; well-defined iridescent display plumage on 
back and sides of neck. Plumage largely dark slate-grey ; wings and tail unmarked. 

C. leucocephala Linnaeus, 1758 
C. squamosa Bonnaterre, 1792 

(b) Speciosa Sub-group 

Feathers of neck and breast white and golden with dark, iridescent edges ; back, 
rump and wing-coverts purplish-chestnut (male) or dark brown (female). Tail 
blackish and unmarked. One species only. 

C. speciosa Gmelin, 1789 

(c) Picazuro Sub-group 

Outer wing-coverts edged white ; mantle and wing-coverts brown, tail with ill- 
defined terminal dark band. 



{ 



C. picazuro Temminck, 1813 
C. corensis Jacquin, 1784 
C. maculosa Temminck, 1813 



(d) Fasciata Sub-group 

Parti-coloured tail with broad, pale terminal band. Iridescent green display 
plumage on hind-neck. One superspecies. 

{C. fasciata Say, 1823 
C. araucana Lesson, 1827 
C. caribaea Jacquin, 1784 

(e) Rufina Sub-group 

Small size. Crown and nape iridescent ; ill-defined pale terminal band on tail; 
plumage mainly vinous-purple. 

C. rufina Temminck, 1810 



TAXONOMY OF THE GENUS COLUMBA 7 

(f) Flavirostris Sub-group 

No iridescence on head or neck ; plumage largely vinous-purple ; tail dark grey 
without markings. One superspecies. 

C C . flavirostris Wagler, 1831 
< C. oenops Salvin, 1895 
tC inornata Vigors, 1827 

Sub-genus Oenoenas 

Small columbids of central and southern America with dark brown and greyish- 
purple or reddish-purple plumage ; small bills, strongly rounded tails, obsolescent 
display plumage on hind neck and no markings on wings or tail. Consists of only 
four species. It seems purposeless to sub-divide further this small and very homo- 
geneous assemblage into sub-groups, which could only be done, if at all, on the shorter 
tails of two of them, nigrirostris and goodsoni. 



{ 



C. subvinacea (Lawrence, 1868) 
C. plumb ea Vieillot, 1818 
C. nigrirostris Sclater, 1859 (i860) 
C. goodsoni Hartert, 1902 



Sub-genus Turturoena 

Small columbids inhabiting the Ethiopian region. Bright metallic iridescence 
on hind-neck. Marked sexual dimorphism except in the island form malherbii. 
One superspecies. 

CC. delagorguei Delagorgue, 1847 
< C. iriditorques Cassin, 1856 
I^C. malherbii Verreaux & Verreaux, 1851 

Sub-genus Nesoenas 

Large-sized Mauritian columbid with rounded wings (first primary about equal 
in length to sixth) and rufous tail. One species. 

C. mayeri Prevost, 1843 

NOTES ON SPECIES 

C. palumbus 

The Wood Pigeon is considered by Boetticher (1954) to be most closely related 
to torringtoni, elphinstonii, trocaz and junioniae and to be much nearer to fasciata 
and allied American species than to oenas or unicincta. He grouped these, and some 
other pigeons, close together because they all show a high degree of ornamentation 
on the neck feathers, similar relative lengths of wing and tail and arboreal habits. 

Certainly the conspicuous white or cream-coloured patches on its neck give 
palumbus a superficial resemblance to such species as fasciata which also have con- 
spicuous white areas in the display plumage of the neck. In its colour-pattern and 



TAXONOMY OF THE GENUS COLUMBA 




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TAXONOMY OF THE GENUS COLUMBA g 

even in the individual colours, however, it agrees more closely with unicincta and 
oenas. The difference in coloration of the display plumage on the necks of oenas 
and palumbus, due to the white patches of the latter, can be interpreted as an iso- 
lating mechanism (see Sibley, 1957) as can the difference in their advertising calls. 
The bowing display of palumbus is identical with that of oenas and its pre-copulatory 
behaviour is very similar to that of oenas, guinea and livia (see Goodwin, 1956a) 
and differs from that oifasciata as described by Whitman (1919). It is true that in 
some points of structure and deportment palumbus bears more resemblance to 
fasciata than to oenas but this may be attributed to the fact that palumbus and 
fasciata are largely arboreal and obtain much of their food from the tips of slender 
branches whereas oenas is almost entirely a ground feeder. Geographical distribution 
also seems to favour the linkage of palumbus and oenas. In my opinion, and excepting 
obvious derivatives on the Atlantic islands (see trocaz and bollii), palumbus has 
closest affinities with unicincta and is nearer to oenas than it is to pulchricollis or 
fasciata or to their close allies. At the same time its undoubted resemblance to these 
latter emphasize the difficulties of generically separating the Eurasian and American 
groups of Columba. 

C. trocaz 

The Madeiran Pigeon appears to be a fairly recent derivative of palumbus. In 
its voice and bowing display it closely resembles that species and in appearance 
it differs only in its slightly darker coloration, in lacking the white signal markings 
on the wings and in its slightly longer legs and more sloping forehead. As, however, 
it would seem to have diverged sufficiently not to have interbred with later Wood 
Pigeon invaders, now distinguished racially as C. palumbus maderensis, it has to 
be accorded specific rank. 

C. bollii 

Bolle's Pigeon is considered by Peters (1937) and others to be conspecific with 
trocaz. Both appear to be derivatives of palumbus but evidence suggests that they 
did not share a common ancestry subsequent to their differentiation from the parent 
stock. Although both have lost the white wing patch of palumbus the white neck 
patch has modified in different directions in each. In trocaz it has become a dull silvery 
grey and increased in area whereas in bollii it has become a small glossy brown 
mark. Since trocaz cannot now be considered conspecific with palumbus it seems 
better to give bollii specific status also. 

C. unicincta 

Boetticher considers the Afep Pigeon to be a relatively primitive species, most 
closely related to maculosa, because it lacks display plumage on the neck. It seems 
to me much more likely that loss of display plumage has occurred at a relatively 
late stage in the evolution of both species and does not indicate any close phylo- 
genetic relationship between them. In its general colour pattern unicincta agrees 
fairly closely with palumbus and its tail markings are identical with those of that 



io TAXONOMY OF THE GENUS COLUMBA 

species. Two features in which unicincta differs from palumbus are in its pale-edged 
wing-coverts and in its grey feet and bill. Since, however, these features, like the 
lack of iridescence on the neck, are similar to those in juveniles of palumbus I do 
not think they diminish the likelihood of close phylogenetic relationship of unicincta 
and palumbus, which their other features, including the little that has been recorded 
about the behaviour and ecology of unicincta, seem to indicate. 

C. oenas 

The Stock Dove is usually considered to be much more closely related to livia 
than to palumbus but in fact it forms a connecting link between the two in its taxo- 
nomic characters as well as in its ecology and behaviour (Goodwin, 1956a). Thus it 
agrees with palumbus in the distribution of the display plumage on its neck, in its 
glossless mauve-pink breast and in the form of its bowing display, but its wing and 
tail markings and its proportions are closer to livia. In view of the intermediate 
position of this, the type species of Columba, generic separation of " wood pigeons " 
and " rock pigeons " becomes untenable. 

C. eversmanni 

Eversmann's Stock Dove is extremely close to oenas in appearance, differing only 
in its smaller size, yellow irides and in having the head as well as the breast mauve- 
pink, or with a strong mauve-pink tinge, and in sometimes having a white instead 
of a grey rump. I concur with Peters in thinking that the evidence at present available 
warrants their both being considered full species. Stuart Baker (1928) put eversmanni 
as a race of oenas stating that " intermediate forms are not infrequent ". He gave 
no evidence for this other than mentioning some grey-rumped eversmanni having 
been obtained in Afghanistan. Except in this one feature grey-rumped individuals 
do not resemble oenas any more closely than do white-rumped ones. Stuart Baker's 
further remarks about the affinities of the two show that he knew little about the 
behaviour and ecology of oenas. The breeding range of eversmanni as indicated 
by Dementiev et al. (1951) has a considerable overlap with that of oenas. That they 
retain their respective characteristics where they are, apparently, sympatric, is 
evidence of their specific status. 

C. oliviae 

The Somaliland Pigeon can be regarded as forming a superspecies with eversmanni. 
The differences between them can be explained by the adaptation of oliviae to its 
torrid desert habitat. As well as acquiring a paler and browner coloration it has 
lost the black wing bars, probably quite recently, as many specimens show a few 
blackish flecks where these are in eversmanni. The loss of these signal markings may 
have resulted from a need for more perfect camouflage or simply to lack of selection 
for them subsequent to isolation from other closely related species. Unfortunately 
nothing appears to have been recorded of the behaviour or breeding habits of this 
species. In spite of its alternative vernacular name of " Somali Rock Pigeon ", 
oliviae is no more closely related to livia than are eversmanni or oenas. 



TAXONOMY OF THE GENUS COLUMBA n 

C al orques 

The White-collared Pigeon has a white neck ring like that oifasciata and acuminate 
neck feathers like arquatrix. But as it resembles these species in no other respects 
it is highly unlikely that these likenesses indicate close relationship. Its tail pattern 
and the black markings on its wings seem to indicate relationship to oenas and livia. 
Taibell (1954) describes a bowing display like that of oenas. The lack of strong 
contrast between the black wing markings and their dark grey background, the slight 
degree of iridescence of the neck feathers and the partially concealed white wing 
patch may indicate that albitorques is at a stage of evolution where it is beginning 
to lose some of its display and signal markings, presumably as a result of being 
now ecologically isolated from related species. On the whole the appearance and 
taxonomic characters of this pigeon suggest a rather closer relationship with livia 
and guinea than with oenas and I have, accordingly, placed it in the livia sub-group. 

C guinea 

It is fairly certain that the Speckled Pigeon is a close relative of livia (see Goodwin, 
1956 and 1956a) in spite of their differences of colour. They resemble each other in 
having display plumage on the front as well as on sides and back of the neck and 
in having the feathers of this display plumage bifurcated ; there are also many 
close resemblances of voice and behaviour. On the evidence of the blood antigens 
Cumley & Irwin showed that biochemically guinea is closer to palumbus than it is 
to livia or oenas. This is supported by two points of behaviour — its calls and postures 
immediately after copulation and its methods of fighting. Thus what is known 
of this species emphasizes the essential homogeneity of the oenas species-group in 
spite of the rather considerable, if somewhat superficial, taxonomic and behavioural 
variations within it. I think, however, that in sub-dividing this group guinea should 
be put in the same sub-group as livia. 

C. leuconota 

The Snow Pigeon's plumage and general habits suggest a close relationship with 
livia and rupestris. The great differences in colour of the head and neck between 
leuconota and rupestris is almost certainly to be considered as an isolating mechanism 
and of little importance as an indicator of phylogenetic disparity. The same may 
be true of the apparently considerable difference between the voice and bowing 
display of leuconota (Newman, 1911) and that of livia. The black and white pattern 
on the tail of leuconota appears different from that of livia and other allied species 
but is essentially similar, the differences being due to a reduction of the black terminal 
band and an extension of the black basal area on the outer rectrices. 

C. livia 

The Rock Pigeon gives an impression of standing far apart from palumbus in 
appearance and behaviour. Boetticher (1954) indeed puts them as the terminal 
twigs on the two mutually diverging main branches of his family tree of Columba. 
The differences of form, behaviour and coloration shown by livia are largely due 



12 TAXONOMY OF THE GENUS COLUMBA 

to its adaptations to nesting and sheltering in caves and rock fissures and feeding 
in open tree-less country. In fact I think that livia and palumbus are fundamentally 
much alike and that they represent the extremes in a single rather large species-group. 
Although livia now occurs in some moderately wooded regions it almost certainly 
evolved in a treeless or almost treeless environment. Its points of resemblance 
to rupestris, guinea and oenas are discussed under those species. 

C. rupestris 

The Blue Hill Pigeon is usually accorded specific rank although Austin (1948) 
put it as a race of livia but without giving reasons for this decision. Over most of 
their range the two are allopatric, rupestris replacing livia in the higher and colder 
parts of central and eastern Asia. The differences between them are certainly such 
as suggest racial rather than specific status. Rupestris has a proportionately smaller 
bill and a reduced amount of melanin in the plumage resulting in a white instead 
of a grey central bar in the tail, smaller black wing bars, paler grey ground colour 
and less intensely iridescent display plumage ; all differences such as might be 
expected in a race inhabiting a colder climate. With the possible exception of the 
white tail bar none of these cause as great difference in appearance as exists, for 
example, between C. livia gymnocyclus and C. /. dakhlae or C. I. livia and C. /. 
intermedia. In its habits and ecology (Schafer, 1938) rupestris apparently differs in 
no essentials from livia. 

These facts suggest that the two are conspecific. On the other hand rupestris 
is said to associate with livia in parts of northern India (Stuart Baker, 1913) and 
it is potentially in contact with feral specimens of livia in many parts of its range. 
La Touche (1934) shot a specimen of C. livia intermedia (sic) out of a flock of rupestris 
in eastern China. The voice of rupestris, as described by Salim AH (1949), would 
appear to differ much from that of livia. The evidence is thus conflicting. La Touche 
considers that the occurrence of mixed flocks suggest that rupestris and livia inter- 
breed but this might equally well be evidence to the contrary. If, as is likely, the 
term " mixed flock " has been applied to aggregations of rupestris and (? feral) 
livia at feeding grounds then such associations are of no significance either way. 
There are no specimens from among the many from North India and the Himalayas 
in the National Collection that appear to be hybrids or intergrades between the two. 
It must be admitted, however, that such birds might be difficult to detect, unless 
they were intermediate in coloration of the tail. Until the matter can be investigated 
more conclusively it seems best to accept the majority opinion that rupestris is 
a full species. Since the evidence suggests strongly that, at least as far as genuinely 
wild individuals are concerned, livia and rupestris are allopatric when breeding they 
should, I think, be considered members of a superspecies. 

C. junioniae 

The coloration and size of the Laurel Pigeon suggest relationship to palumbus. 
On the other hand the colour-pattern of its tail is similar to that of albinucha and 
fasciata. Meade-Waldo (1889) described it as having a very distinctive gait when 



TAXONOMY OF THE GENUS COLUMBA 



13 



walking on the ground and a " soft, flopping flight •" which was unlike that of any 
other pigeon known to him. He found that on the Island of Gomera, in the Canaries, 
it kept mainly to scrub-covered slopes at relatively low altitudes, being replaced 
in the high mountain forest by C. bollii. It lays one egg to a clutch. The Laurel 
Pigeon seems most likely to be either an offshoot from primitive palearctic Columba 
stock prior to its subsequent speciation or an early offshoot from palumbus. It is 
impossible to do more than guess at its affinities from an examination of skins and 
the little that has been recorded of its behaviour. A detailed comparison of its 
anatomical and behavioural characters with those of, at least, palumbus, fasciata 
and albinucha will probably be necessary before its true relationships can be ascer- 
tained. 

C. arquatrix 

Nominate arquatrix with a purple head, silver-grey nape and yellow bill and legs 






KEY TO SHADING 


iiiiii 1 
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Black or dark slate-grey 

Dark purple or dark 
purplish-chestnut 

Greyish-white or silver-grey 

Dark purple or purplish-chestnut 
with small white spots 

Dark purple with large pale grey 
spots 

Dark brown and purplish-brown 

Lanceolate display plumage with dark 
bases and shining tips to feathers 






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Fig. 4. Diagrammatic sketches to show colours of the Arquatrix sub-group. Left to 
right : a. thomensis, a. sjostedti, a. arquatrix and (bottom) hodgsoni, polleni and albinucha. 



has always been considered, rightly in my opinion, conspecific with sjostedti from 
the Cameroon Mountains, which has an entirely dark grey head, red orange-tipped 
bill and dark purplish legs and feet. The Sao Thome representative, thomensis, is, 
however, currently given specific rank. It should, I think, be put as a race of arqua- 
trix. In plumage it is just a dull version of the latter with the spotting on wing- 
coverts and undersides and the lanceolate display plumage on the neck showing, 
apparently, the beginning of that obsolescence of markings that so often takes place in 
island forms. It otherwise shows characters of both mainland forms, having an 



i 4 TAXONOMY OF THE GENUS COLUMBA 

entirely dark grey head like a. sjostedti but yellow legs and bill like a. arquatrix. It 
has a somewhat longer tail than either mainland form but this alone does not, 
in my opinion, warrant its specific separation from them. 

C. polleni* 

In spite of its almost uniform dark brown plumage polleni is also a representative 
of arquatrix. Although, presumably in the course of long isolation on the Comoro 
Islands, it has lost the rich colours of arquatrix it still retains lanceolate display 
plumage on the hind neck and the yellow legs and bill. Of three skins in the National 
Collection one shows faint pale spots on the underparts exactly similar to, though 
less pronounced than the spots of arquatrix. It has, however, reached a degree of 
differentiation from arquatrix that justifies, in my opinion, its specific rank, though 
it should be considered as forming a superspecies with arquatrix and hodgsoni. 

C. hodgsoni 

Verheyen (1955) considers this Indian species conspecific with arquatrix. That 
they are geographical representatives of the same stock cannot be doubted but 
hodgsoni differs from arquatrix in several minor points of coloration of plumage and 
soft parts and in its much greater sexual dimorphism. In view of this and of their 
wide geographic separation I think it is better to give them specific rank as members 
of the same superspecies. 

C. albinucha 

In coloration, colour-pattern and display plumage the White-naped Pigeon very 
closely resembles arquatrix. It differs in having red bill and feet, a patterned tail 
with conspicuous pale terminal bar and in lacking the white spots on the wing- 
coverts, although its underparts are spotted in the same manner as those of arquatrix. 
It is also a little smaller. The two are sympatric but their ecology and habitat 
preferences differ (Van Someren, 1949). Nothing seems to have been recorded of the 
displays, nesting or clutch-size of albinucha. Although there can be no doubt that 
they should be regarded as good species, the resemblances between albinucha and 
arquatrix are almost certainly due to close phylogenetic affinity and not to conver- 
gence. 

C. elphinstonii, C. pulchricollis and C. torringtoni 

Although differing in size and coloration these three forms all resemble each other 
in colour-pattern, form of display plumage and (so far as is recorded) in habits and 
ecology. It is difficult to decide whether it is best to treat them as races of a single 

* Through the kindness of Mr. C. W. Benson I have recently been able to examine the fine series of polleni 
collected on the B. O. U. Centenary Expedition to the Comoro Islands. These specimens show that 
the unsexed skins I had previously seen were all females and all somewhat "foxed". In fact males are a 
brownish purple on head, upper mantle and breast and the dark brown parts of both sexes are suffused 
with grey, especially on the rump. They thus resemble arquatrix more closely than I had thought and 
Mr. Benson informs me that the calls of the two are similar ; possibly identical. I, therefore, now consider 
that polleni should be treated as a race of arquatrix and not as a separate species. 



TAXONOMY OF THE GENUS COLUMBA 15 

species or as members of a superspecies. Their ranges nowhere overlap and they 
have, presumably, been long isolated from contact with each other. As they have 
always been accorded specific rank it seems to me best to regard them as members 
of a superspecies. 

C. punicea 

The Purple Wood Pigeon most closely resembles argentina in colour-pattern 
although its purplish-chestnut body colour gives it a very different superficial 
appearance. This species is said sometimes to lay one and sometimes two eggs per 
clutch (Stuart Baker, 1913). If this is really the case — no one other than Stuart 
Baker appears to have found nests with more than one egg — this is an interesting 
intermediate condition between the species laying one egg and those laying two. 

C. argentina 

The Silver Pigeon shows a striking resemblance to the Pied Imperial Pigeons in 
colour and colour-pattern. Boetticher (1954) considered it more closely related to 





Fig. 5. Diagrammatic sketches to show colour-patterns of tails and outermost tail 
feathers of left, Ducula bicolor; right, Columba argentina. 

Ducula than to Columba. I have been unable to find any description of the anatomy 
of this pigeon which would show whether its stomach and intestines are of Ducula 
or Columba type. From its plumage characters, I think, however, that it is rightly 
placed in Columba. It shows the same type of sexual dimorphism — female a slightly 
duller and browner grey — in plumage colour as do other grey Columba species. It 
resembles punicea in the colour-pattern of its wings although the coloration of the 
two is different. Superficially the tail appears to be similarly marked to that of the 
Pied Imperial Pigeon but the pattern on the outer retrices (see Text-fig. 5) is quite 
different. I think, therefore, that argentina' s resemblance to the pied Ducula species 
is due to convergence and that it is best regarded as a rather aberrantly coloured 
member of the Elphinstonii species-group. It is another example of a Columba 
species which appears to have lost the display plumage on the neck, a point compatible 
with its presumed affinity to punicea which appears to show a regressional stage of 
this display plumage. 

C. palumboides 

In appearance, at least of the skin, the Andaman Wood Pigeon suggests a link 
between the Elphinstonii and Janthina species-groups. It seems likely to be nearer 



16 TAXONOMY OF THE GENUS COLUMBA 

to the former on geographical grounds. Hume (1867) who collected some specimens 
and had a captive bird under observation was impressed by its " mode of holding 
itself and its broad substantial body " which he considered argued relationship to 
Ducula rather than to Columba. This may not be of importance as in so far as it is 
adapted to an arboreal life ; any large pigeon is bound to show some resemblance 
to Ducula as indeed does the common Wood Pigeon palumbus. A comparison of 
the alimentary tracts and a comprehensive study of the foods taken by this and 
other very arboreal Columba species with those of the Ducula species living in the 
same regions would be of great value. 

C. janthina 

The Black or Japanese Wood Pigeon and C. vitiensis, seem best considered as 
members of a single superspecies. C. j. nitens with its distinctive purple-brown 
head is somewhat intermediate, but in most respects is closer to nominate janthina 
than to any race of vitiensis. Stresemann (1939) suggests New Guinea as the place 
of origin of these forms and gives a map of their distribution and the probable 
directions of their former spread from the Papuan regions. 

C. vitiensis 

Although generally arboreal, so far as is known, C.v. halmaheira has been recorded 
nesting on the ground among thick cover (Warner, 1949) and Mr. I. C. J. Galbraith 
informs me that the natives on the Solomons told him that ground nesting was not 
infrequent in this species. In its bowing display (Newman, 1910) the bird moves rather 
slowly and does not erect its tail. 

C. norfolciensis 

This species appears to be a derivative of vitiensis. Males differ strikingly from 
the parent species in having the head and underparts white, or, in new plumage, 
creamy buff with a pink tinge. Females, however, have these parts whitish-grey 
or greyish-buff with some iridescence on the crown of the head and the pale throat — 
as in vitiensis — almost pure white and fairly clearly demarcated. Whether norfolciensis 
merits specific recognition is arguable but in view of the great difference in the 
appearance of the males, which makes it highly likely that if at some future period 
it were to invade the range of vitiensis or vice versa the two would not interbreed, 
and the nomenclatorial changes that would be involved, it seems preferable to accord 
it specific rank within the same superspecies. 

C. versicolor 

This extinct species appears to have been a very close relative of janthina. Kuroda 
(1925) considers it a race of janthina. Both forms occurred, however, on the Bonin 
Islands and I think therefore that Peters (1937) was right to give them specific rank 
in spite of their obvious affinities in form and coloration. 



TAXONOMY OF THE GENUS COLUMBA 



17 




18 TAXONOMY OF THE GENUS COLUMBA 

C. jouyi 

Kuroda (1925) considers jouyi conspecific with janthina. Its distribution as given 
by him overlaps with nominate janthina on Okinawa. Jahn (1940) does not, however, 
record jouyi from Okinawa. Besides the large silver patch on its hind-neck and 
mantle — which presumably functions as a recognition mark and probably also as 
an isolating mechanism — jouyi is a larger, longer-tailed bird and shows only slight 
traces of iridescence other than on the neck and nape. I therefore think it best to 
follow Peters in giving jouyi specific status. 

C. pallidiceps 

The Yellow-legged Pigeon appears to be a derivative from vitiensis stock. It 
now overlaps C. vitiensis halmaheira on some of the Solomon Islands (Mayr, 1934) 
although only pallidiceps occurs in New Britain, where it perhaps originated. Al- 
though much alike in general appearance the two differ strikingly in the coloration 
of the head, silver-grey in pallidiceps, dark purple with a white throat in vitiensis; 
and in the feet, which are red in vitiensis and yellow in pallidiceps. The coloration of 
pallidiceps is suggestive of that of norfolciensis (q.v.) whose close relationship to 
vitiensis is beyond doubt. It would be interesting to know if the difference in leg 
colour between the two serves in any way as an isolating mechanism. It would 
seem rather unlikely particularly as it is not correlated with any difference in bill 
colour, which is mainly red in both species. In other pairs of Columba species (e.g. 
arquatrix and albinucha) in which one has yellow and one red legs there is a correlated 
difference in bill colour. 

C. leucocephala 

The White-crowned Pigeon is very closely related to the Red-necked Pigeon 
C. squamosa. Both have the same highly-developed and beautifully (but differently) 
coloured display plumage on the back and sides of the neck, are of similar size and 
inhabit the same geographical areas. They may, however, be ecologically isolated 
as in Cuba (Barbour, 1943) ; leucocephala is a coastal and squamosa a highland 
form. The striking difference in coloration of the head, eyes and display plumage 
in these otherwise almost identical forms suggests that these differences function 
or have functioned as isolating mechanisms. 

It seems probable that this pigeon lays but one egg to a clutch (Wetmore and 
Swales, 1 931) in spite of an earlier statement (Gundlach, 1874) that it lays two. 

C. squamosa 

The close relationship between the Red-necked and White-crowned pigeons 
has been discussed under leucocephala. The sexual dimorphism in squamosa is much 
less than in leucocephala but readily apparent when skins in a like state of plumage 
are compared. Probably this pigeon also lays only one egg (Wetmore, 1927) in 
spite of earlier statements (Gundlach, 1874) to the contrary. 



TAXONOMY OF THE GENUS COLUMBA 19 

C. speciosa 

The Scaled Pigeon appears to be fairly close to squamosa and leucocephala. In 
some respects its plumage is intermediate between that of these species and of 
picazuro but I do not agree with Verheyen that speciosa and picazuro should be 
generically separated from squamosa and leucocephala. In speciosa the dark edgings 
to the feathers are found throughout most of its plumage but are intensified in 
contrast and brilliance on the neck, thus giving a suggestion of the condition obtaining 
in leucocephala and squamosa. The Scaled Pigeon shows strong sexual dimorphism, 
the male having the mantle and wing-coverts rich purple (fading to chestnut) whereas 
those of the female are dull brown. 

C. picazuro 

The Picazuro Pigeon has the display plumage on the neck less highly developed 
than in leucocephala and squamosa but of an essentially similar kind. According to 
Hudson (1920) its habits, gait and behaviour are similar to those of palumbus but 
except for a vivid contrast of its gait on the ground with that of maculosa he gives 
no details. Hudson said that it lays two eggs, Venturi (see Hartert, 1909) that 
sometimes one and sometimes two eggs are laid. Wetmore (1926) found a nest with 
one egg. He also said that the display flight is like that of livia, a rather surprising 
fact in an essentially woodland species. 

C. corensis 

The Bare-eyed Pigeon can be considered as forming a superspecies together with 
picazuro. Its main differences, the much paler coloration and the very extensive 
amount of bare orbital skin, are probably adaptations to the hot, coastal areas 
where it lives. 

C. maculosa 

I think the Spotted Pigeon is most closely related to picazuro. It has evidently 
become more fully adapted to feeding on the ground in open country. Its resemblance 
in wing /tail proportions and gait to livia is almost certainly due to convergence in 
feeding habits. This is another species which has no display plumage on the neck, 
probably having lost it at a relatively recent date. If so, no close relationship to 
other species sharing this negative character is indicated. 

C. fasciata 

The Band-tailed Pigeon is reminiscent of C. arquatrix in its yellow bill and feet. 
The pattern of the bill is unusual in adult Columba but very much like that of Columba 
nestlings, only the dark sub-terminal band having " spread " to the tip of the bill and 
the lighter base turned bright yellow. The copulation ceremony of this species 
(Whitman, 1919) evidently differs in several details from that of European Columba 
species. 



20 TAXONOMY OF THE GENUS COLUMBA 

C. araucana 

Although the Chilean Pigeon is very closely related to the southern form of the 
Band-tailed C '. fasciata albilinea it has, I think, diverged sufficiently to be given specific 
rank. Besides the plumage differences (purplish-maroon on mantle and scapulars) 
it has a dark bill and red legs. It can be considered as forming a superspecies together 
with fasciata and caribaea. 

C. caribaea 

The Jamaican Band-tailed Pigeon is clearly derived either from fasciata, araucana 
or from some comparatively recent form ancestral to them both. In plumage it 
differs from fasciata only in being paler, in having lost the white neck-ring and having 
less well developed iridescent display plumage on the neck. It differs also in having, 
like araucana, a dark bill and red legs. 

C. cayennensis 

The Rufous Pigeon is intermediate in many respects between the members of 
the " fasciata " superspecies and those of the superspecies " flavirostris ". It resembles 
the former in the colour-pattern of its tail feathers and in having display plumage 
on the hind neck but in its other plumage characters it is closer to flavirostris. Little 
or nothing seems to be recorded about its breeding habits in the wild but two clutches 
of one egg each were laid by captive specimens (Newman, 1910). 

C. flavirostris 

The Red-billed Pigeon forms a superspecies together with oenops and inornata. 
All three completely lack display plumage on the neck thus presenting a superficial 
resemblance to the Oenoenas species. 

C. oenops 

This is the Peruvian representative of flavirostris. In appearance it differs only 
in the colour of its bill, which is yellow, or yellow tipped with black, instead of pink 
and whitish. Also the males have a larger area of ruddy purple on the plumage of 
the upper parts but females and juveniles are intermediate in this respect between 
males of their own species and flavirostris. 

C. inornata 

This island representative of flavirostris differs only in its larger size, black bill 
and grey, instead of red, orbital skin. 

The Sub-genus Oenoenas 

Salvadori (1893) characterized this sub-genus as having " general plumage, 
brown, more or less vinous, and with no light spots on the neck, but sometimes with 
concealed reddish spots, more or less obsolete, on the hind-neck ". Actually the 
" reddish spots " are appreciably lighter than the adjacent areas of the feathers 



TAXONOMY OF THE GENUS COLUMBA 21 

that show them and these pigeons also have, as pointed out by Ridgway (1916), 
rather small bills and strongly rounded tails. Ridgway gave them, and others less 
deserving, full generic rank but subsequent authorities (Peters, 1937 ; Hellmayr 
& Conover, 1942 ; de Schaunsee, 1948) put Oenoenas into the synonymy of Columba. 

The four species concerned, plumbea, subvinacea, nigrirostris and goodsoni are 
all closely related and are more sharply differentiated than any others within Columba, 
mayeri excepted. I do not concur with Boetticher in considering them to be 
closely related to caribaea or that their dull concolorous plumage represents 
the retention of a primitive condition. The Oenoenas species differ from other Ameri- 
can pigeons (except for the quite differently coloured speciosa) in lacking grey on 
the rump, which is concolorous with the mantle and wings or nearly so. The more 
or less obsolescent pale or bronzy-red markings on the feathers of the hind-neck 
are more prominent in females than in males, which suggests that so far from repre- 
senting display plumage in process of acquisition, as Boetticher suggests, the reverse 
is the case. The obsolescent display plumage and general dull coloration of Oenoenas 
seems to be, in the evolutionary sense, a relatively recent development, not the 
retention of primitive characters. 

Oenoenas is most probably an early offshoot from the other American Columba 
that has diverged considerably from the parent stock. Its sombre, concolorous 
dress with complete lack of signal markings and obsolescent display plumage pre- 
sumably evolved as a consequence of increased predator pressure consequent on 
smaller size and /or increased reliance on vocal differences as isolating mechanisms. 

C. plumbea and C. subvinacea 

Only the fact that these pigeons appear to be sympatric in parts of their range, 
makes it possible — on present knowledge — to consider them as specifically distinct. 
Whilst the slight differences of colour and bill size make it possible to identify most 
specimens, it is difficult to believe that they could be sufficient to act as isolating 
mechanisms for the birds themselves. Presumably vocal and behavioural differences 
exist, but they do not appear to have been recorded. A few specimens in the National 
Collection are intermediate in character between plumbea and subvinacea but it is 
impossible to say whether these are hybrids or merely individuals whose slight 
aberrancy is towards the characters of the other species. 

C. nigrirostris and C. goodsoni 

These can certainly be regarded as forming a single superspecies. They may well 
be conspecific but since their taxonomic differences are at least as great (or one 
might better say no less) than those of plumbea and subvinacea, it seems better to 
grant them specific rank until more is known about them. 

The Sub-genus Turturoena 

This is comprised of three allopatric species delagorguei, iriditorques and malherbei 
which replace one another geographically and can be considered as forming a single 
superspecies. Salvadori (1893) separated Turturoena generically from Columba 



22 TAXONOMY OF THE GENUS COLUMBA 

on its smaller size, hind neck with bright metallic colours and the sexes being often 
very dissimilar. Subsequent authorities have disagreed about its status, Peters 
(1937) placing it in the synonymy of Columba and Cave & Macdonald (1955) and 
Mackworth-Praed & Grant (1952) giving it full generic rank. I think, personally, 
that the differences do not merit more than sub-generic rank within Columba. 

Boetticher considers that Turturoena is more closely related to Aplopelia than to 
Columba. Certainly all three species have a similar distribution of iridescence to 
that of the Aplopelia species (but one found also in some species of Columba, and 
other pigeons) and a similar degree and type of sexual dimorphism. However, in 
my opinion, they show more affinities to the Columba species, not only in their 
general appearance but also in their colour-patterns which much resemble those 
found in palumbus and allied species. Mr. R. E. Moreau informs me (in litt.) that 
the cooing of delagorguei sharpei in Tanganyika reminded him very much of that 
of the Wood Pigeon, palumbus. 

The Sub-genus Nesoenas 

Salvadori (1893) placed the Mauritius Pigeon, C. mayeri, in the monotypic genus 
Nesoenas because of its more rounded wing, with the first primary about equal in 
length to the sixth, and its rufous tail. In coloration this pigeon does not closely 
resemble any other Columba but its colour-pattern shows some resemblance to the 
oenas species-group, from which it may possibly be an offshoot. Fortunately this 
species is not extinct as was at one time feared so it is possible that studies of it 
that will serve to indicate its affinities may yet be made. In the meantime I think 
it is better to put Nesoenas as a sub-genus within Columba than to afford it generic 
rank. 

SUMMARY 

This paper revises the genus Columba. The taxonomic and behavioural characters 
of the species are discussed in reference to their probable affinities. 

The absence or obsolescence of the display plumage on the neck is never a primitive 
character in Columba but one that has evolved independently in several species. 

Oenoenas, Turturoena and Nesoenas are given sub-generic rank. It does not seem 
feasible to further sub-divide the genus. There are indications that the American 
species may prove separable on anatomical and biochemical criteria although 
they cannot be separated on external characters. 

REFERENCES 

Ali, S. 1949. Indian Hill Birds. Oxford (p. 173). 

Austin, O. L. 1948. The Birds of Korea. Bull. Mus. Comp. Zool. 101 (1) : 1-130. 
Baker, E. C. Stuart. 1913. Indian Pigeons and Doves. London. 
Barbour, E. 1943. Cuban Ornithology (Mem. Nuttall. Orn. Club. no. 9, pp. 70-72). 
Boetticher, H. von. 1954. Die Taubengattung Columba L. Zool. Anzeiger, 153, 3-4 : 49-64. 
Cain, A. J. 1954. Subdivisions of the genus Ptilinopus. Bull. Brit. Mus. (Nat. Hist.) Zool. 2, 
No. 8. 



TAXONOMY OF THE GENUS COLUMBA 



^3 



Cave, F. O. & Macdonald, J. D. 1955. Birds of the Sudan. London. 

Cumley, R. W. & Irwin, M. R. 1944. The correlation between antigenic composition and 

geographic range in the Old and New World of some species of Columba. Amer. Nat. 

LXVIII, No. 776 : 238-256. 
Dementiev, G. P. et al. 1951. Birds of the Soviet Union, 11 : 6-32. 
Goodwin, D. 1956. Remarks on the Rock and Speckled Pigeons. Bull. Brit. Orn. Club, 76 : 6-9. 

1956a. Observations on the voice and displays of certain pigeons. Avic. Mag. 62 : 17-33 

and 62- 70. 

Gundlach, J. 1874. Neue Beitrage zur Ornithologie Cubas. Journ. Orn. 1874 : 289. 
Hartert, E. & Venturi, S. 1909. Notes sur les oiseaux de la Republique Argentine. Novit. 

Zool. XVI, No. 2 : 260-261. 
Hellmayr, C. E. & Conover, B. 1942. Catalogue of Birds of the Americas, pt. 1, no. 1, Zool. 

Ser. Field Mus. Nat. Hist. XIII, pt. 1, No. 1. 
Hudson, W. H. 1920. Birds of La Plata. Vol. 2, pp. 154-155. 

Jahn, H. 1942. Zur Oekologie und Biologie der Vogel Japans. Journ. Orn. 90, 1-2 : 7-301. 
La Touche, J. D. D. 1934. A handbook of the birds of Eastern China. Vol. 2, p. 209. London. 
Kuroda, N. 1925. A contribution to the knowledge of the avifauna of the Riu Kiu Islands and 

the vicinity. Tokyo. 
Mackworth Praed, C. W. & Grant, C. H. B. 1952. The African handbook of Birds : Ser. 1, 

vol. 1, p. 468. 
Mayr, E. 1931. The birds of Malaita Island. Amer. Mus. Novit. 504, No. nth, 1931, p. 10. 
1934- Notes on some birds from New Britain, Bismarck Archipelago. Ibid. 709, 4th April, 

I934» PP- 4-5- 
Meade-Waldo, E. G. 1889. Notes on some birds of the Canary Islands. Ibis, 6th Ser. No. 1, 

pp. 6-9 and No. 4, pp. 509-510. 
Miculicz-Radecki, M. von. 1949. Betrachtungen zur Stammesgeschichte der Wildtauben. 

Verhand. Deutsch. Zool., Aug. 1949, pp. 55-66. 
Newman, T. H. 1910. Nesting of the White-throated Pigeon. Avic. Mag. 3rd Ser., vol. 1, 

No. 5, pp. 158-164 and i93-!95- 

191 1. The Snow Pigeon. Ibid. Vol. 2, No. 6, pp. 173-178. 

Peters, J. L. 1937. Check-list of the Birds of the World. Vol. 3. Camb. Mass. U.S.A. 
Pollen, P. L. & Van Dam, D. C. 1868. Recherches sur la Faune de Madagascar et ser Dependences. 

Vol. 2, pp. 1 1 2-1 13. Leiden. 
Salvadori, T. 1893. Catalogue of Birds in the British Museum. Vol. 21. 
Schafer, E. 1938. Ornithologische Ergebnisse zweier Forschungsreisen nach Tibet. Journ. f. 

Orn. Supplement, May 1938. 
Schaunsee, R. M. de. 1948. The Birds of the Republic of Columbia, part i. Bogota. 
Sibley, C. G. 1957. The evolutionary and taxonomic significance of sexual dimorphism and 

hybridisation in birds. Condor 59 : 3 : pp. 166-191. 
Taibell, A. M. 1954. Notizie sulla riproduzione in cattivata del Colombo dal collare bianco 

[Columba albitorques) . Rivista It. Orn., 1954 (Seconda serie), pp. 195-203. 
Van Someren, V. G. L. 1949. The birds of Bwamba. Special supplement to the Uganda Journal, 

Vol. 13, pp. 24-25. 
Verheyen, R. 1955. Le Pigeon Bleu [Columba arquatrix Temm.) du Ruwenzori. Le Gerfaut, 

1955. vol. 2, pp. 127-145. 

1957- Analyse du potential morphologique et project de classification des columbiformes 

(Wetmore, 1934). Bui. Inst. Roy. Sci. Bel. Vol. 33, No. 3. 

Warner, D. W. 1949. The White-throated Pigeon nesting on the ground in New Caledonia. 

Auk. 66 : 90-91. 
Wetmore, A. 1927. The birds of Porto Rico, pp. 390-391. 
Wetmore, A. & Swales, B. H. 1931. The birds of Haiti and the Dominican Republic. U.S. 

Nat. Mus. Bull. 155 : 184. 
Whitman, C. O. 1919. The behaviour of pigeons. Posthumous works of C. O. Whitman. 

Vol. 3 : pp. 56-57. Carnegie Inst., Wash. 
zool. 6,i. 2/^ ""*\ 



A REVISION OF THE PLATYSEIINAE 
(MESOSTIGMATA : ACEOSEJIDAE) 

BASED ON MATERIAL IN THE COLLECTIONS 

OF THE 
BRITISH MUSEUM (NATURAL HISTORY) 



G. OWEN EVANS and K. H. HYATT 




BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ZOOLOGY Vol. 6 No. 2 

LONDON: i960 



A REVISION OF THE PLATYSEIINAE 

(ME SO STIGMATA : ACEOSEJIDAE) BASED ON 

MATERIAL IN THE COLLECTIONS OF THE 

BRITISH MUSEUM (NATURAL HISTORY; 



BY 

G. OWEN EVANS and K. H. HYATT 



Department of Zoology, 
British Museum (Natural History) 



Pp. 25-101, 204 Text- figures 




BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY Vol. 6, No. 2 

LONDON: i960 



THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical Series. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

This paper is Vol. 6, No. 2 of the Zoological 
series. 



Trustees of the British Museum, i960 



PRINTED BY ORDER OF THE TRUSTEES OF 
THE BRITISH MUSEUM 

Issued February, i960 Price Twenty-four Shillings 



A REVISION OF THE PLATYSEIINAE 

(MESOSTIGMATA : ACEOSEJIDAE) BASED ON 

MATERIAL IN THE COLLECTIONS OF THE 

BRITISH MUSEUM (NATURAL HISTORY) 

By G. OWEN EVANS & K. H. HYATT 

CONTENTS 

Page 

Introduction ........... 27 

External Morphology ......... 28 

Classification ........... 33 

Genus Platyseius Berlese ........ 34 

Genus Plesiosejus gen. nov. ........ 42 

Genus Sejus C. L. Koch ........ 49 

Genus Zerconopsis Hull . . . . . . . . 91 

Summary ............ 100 

References ........... 100 

INTRODUCTION 

The family Aceosejidae is divided into two subfamilies, Aceosejinae and the Platy- 
seiinae, on the basis of the chaetotaxy of the gnathosoma and the tarsi of legs II-IV 
(Evans, 1957). The classification of the Aceosejinae has been outlined by Evans 
(1958) but no revisionary work has yet been undertaken on the Platyseiinae which, 
at present, contains six genera, namely, Sejus C. L. Koch, 1843, Cheiroseius Berlese, 
1916, Platyseius Berlese, 1916, Episeius Hull, 1918, Zerconopsis Hull, 1918, and 
Episeiella Willmann, 1938. 

Koch (1836) originally assigned a number of species to the genus Sejus but did 
not designate the type species of the genus, Sejus viduus C. L. Koch, until 1843. 
Berlese (1892) accepted Koch's wide concept of the genus and in 1913 proposed 
the family Seiidae for its reception together with eight other genera. Later in 1916, 
the same author revised his concept of the Seiidae and stated that Seius (= Sejus), 
with Sejus togatus C. L. Koch as the t}^pe, was not confamilial with the other genera 
he had included in the family in 1913. Thus, the family Seiidae was reserved for 
those species having the general characteristics of Sejus togatus. Subsequent 
workers (Vitzthum and Tragardh) accepted Berlese's definition of Sejus and the 
family became established in the literature until 1936 when Oudemans noted the 
error in considering 5. togatus to be the type of Sejus. Consequently Sejus Berlese 
nee Koch became a synonym of Liroaspis Banks in spite of Tragardh's remark that 
" this incident offers yet another strong argument in favour of establishing nomina 
conservanda" (Tragardh, 1946). We are following Sellnick (see Willmann, 1953) 

ZOOL, 6. NO. 2. 3 



28 



THE PLATYSEIINAE (MESOSTIGM AT A : ACEOSEJIDAE 



who considers Epicrius corniger Berlese (1891) to be a synonym of Sejus viduus C. L. 
Koch. According to our generic concept Cheirosetus, Episeius and Episeiella are 
synonymous with Sejus. 

The present revision of the Platyseiinae is based on material in the Collections of 
the British Museum (Natural History). The major collections were made by the 
late A. H. G. Alston (Indonesia), P. F. Bellinger (Jamaica), Miss T. Clay (Sikkim), 
G. Owen Evans (Uganda), K. H. Hyatt (Nepal), P. Wygodzynsky (Argentina) and 
M. E. Bacchus, P. N. Lawrence and J. T. Salmon (British Isles). The type material, 
unless otherwise stated, is deposited in the British Museum (Natural History). 



EXTERNAL MORPHOLOGY 

Gnathosoma 

The chelicerae are chelate-dentate in all postembryonic developmental stages and 
show a marked uniformity in shape and dentition throughout the subfamily. The 






Fig. 1-3. Chaetotaxy of the venter of the gnathosoma and palptrochanter, femur and 
genu in the larva (1), protonymph (2), and deutonymph (3) of Plesiosejus italicus 
(Berlese). 

1, rostral setae ; 2, external posterior rostral setae ; 3, internal posterior rostral setae ; 
4, capitular setae ; n2, setae appearing in the deutonymph. 



THE PLATYSEIINAE (MESOSTI GM AT A : ACEOSEJIDAE) 29 







25 




Figs. 4-9. Sejus necorniger (Oudemans), dorsum of larva (4), protonymph (5), and 

deutonymph (6). 
Plesiosejus italicus (Berlese), dorsum of larva (7), protonymph (8), and deutonymph (9). 



3 o THE PLATYSEIINAE (ME SOSTI GM AT A : ACEOSEJIDAE) 

spermatophoral process on the movable digit in the male shows considerable inter- 
specific variation in form (e.g. Text-figs. 33 and 41) and appears to be a useful 
taxonomic character. In all stages the processes from the arthrodial membrane 
at the base of the movable digit are simple and setiform. 

The postembryonic development of the chaetotaxy of the venter of the gnathosoma 
and of the three basal movable segments of the pedipalp is typically parasitoid 
(Text-figs. 1-3). The rostral setae (1) in the immature stages and in the adults 
are long and usually whip-like as are the internal palptrochanter setae which appear 
in the deutonymph. (This characteristic structure of these two pairs of setae is a 
useful character for separating the platyseiines from the closely related aceosejines). 
The corniculi are simple and the specialized seta at the inner basal angle of the palptarsus 
is two pronged. With the exception of Sejus clayi sp. nov., the tectum is basically 
tridentate with the processes subequal in length and inconspicuously divided distally 
(Text-figs. 28, 32 and 76). In S. clayi the median process is extremely long and Y- 
shaped with the lateral processes relatively shorter (Text-fig. 117). 



Idiosoma 

The idiosoma is usually elongate-oval in outline although in Platyseius it may be 
subcircular. 

Dorsum : the dorsum of the larva has two weakly sclerotized shields. In Sejus 
and Plesiosejus the anterior shield bears nine pairs of setae and the pygidial four 
pairs 1 (Text-figs. 4 and 7). The striated cuticle between these sclerotized areas 
carries two pairs of J setae (Ji and J2), three pairs of Z setae (Z1-Z3) and one 
pair of S setae (? S3). Three or more platelets are also present on the cuticle. We 
have seen three main types of protonymph which chiefly differ in the chaetotaxy of 
the posterior half of the dorsum. All have an anterior shield with eleven or twelve 
pairs of setae and a pygidial shield. In the Sejus-type (Text-fig. 5) there are five 
pairs of setae in the J series ; Ji and J2 being situated on the cuticle between the 
anterior and pygidial shields. The protonymphs of both Platyseius and Plesioseius, 
on the other hand show a deficiency in the chaetotaxy of the J series. The former 
have only two pairs in the J series (J4 and J5) and the latter three pairs (J3-J5). 
There are no setae of the J series present on the cuticle between the anterior and 
pygidial shields in these genera (Text-fig. 8). All the deutonymphs we have 
examined have a laterally incised dorsal shield which almost entirely covers the 
dorsum of the mite. The chaetotaxy of the " anterior dorsal shield " (the region 
anterior to the incisions) is relatively constant but the number of setae on the 
"posterior shield" which is largely determined in the protonymph shows con- 
siderable variation. Four types may be readily recognized : 

1. Sejus-type : J, Z and S series each with five pairs of setae (Text-fig. 6). 

1 The system of nomenclature for the dorsal chaetotaxy follows that proposed by Sellnick (1944) and 
added to by Hirschmann (1957). The latter author has used one system of nomenclature throughout 
the suborder and by so doing has attempted to homologize setae in widely differing groups. This may 
be possible in the larval stage but becomes increasingly difficult and purely subjective in subsequent 
developmental stages because of the increased number of setae, especially in those forms exhibiting 
hypertrichy. 



THE PLATYSEIINAE (ME SOSTIGM AT A : ACEOSEJIDAE) 



3i 



2. Zerconop sis-type : J and Z series each with five pairs, S series with four 
pairs. 

3. Platyseius-type : J series with two pairs (J4 and J5), Z and S series each 
with five pairs. 

4. Plesiosejus-type : J series with three pairs (J3-J5), Z and S series each 
with five pairs (Text-fig. 9). 

The dorsal shield in the adults is entire except in S. clayi, Zerconopsis labradorensis 
sp. nov. and Zerconopsis muestari (Schweizer) which retain the deutonymphal 
incisions and Sejus ornatus sp. nov. which has wide incisions between S2 and S3 
(Text-fig. 168). The chaetotaxy of the " posterior dorsal shield " is essentially the 
same as in the deutonymph so that the above four groups may also be recognized 
in the adult. The surface of the shield is invariably reticulated and the majority 




10 11 

Figs, io-ii. Dorsal shield of ^PUsiosejus italicus (Berlese). Fig. 10, lateral view. Fig. n, 

transverse section. 



of the species have a median longitudinal ridge (Text-figs. 10 and 11). The position 
of the highest point along this ridge varies throughout the group. It is usually 
located in the region of setae I4-J2 but in some species it may appear as a distinct 
tubercle between J4 and J5. The extreme attenuation of the posterior margin of 
the anterior shield in the protonymph of Plesiosejus italicus (Berl.) may be due to 
the development of the median ridge (cf. Text-figs. 8 and 36). 

Venter : The tritosternum is well developed in all stages and in the adult comprises 
a long slender base and a pair of strongly pilose laciniae. Pre-endopodal shields 
may be present in some species. 

The development of the chaetotaxy of the intercoxal region is normal ; the larva 
having three pairs (homologous with sternal setae I— III in the adult), the proto- 
nymph four pairs (homologous with sternal setae I— III and the genital setae in the 
adult) and the deutonymph five pairs (the extra setae appearing at this stage being 
homologous with the metasternals in the adult). The female has a strongly sclero- 
tized sternal shield bearing three pairs of setae and the metasternals are normally 
situated on discrete platelets. 

The genital shield is wedge shaped and except in S. clayi and certain species of 
Zerconopsis has a pair of setae. A sterniti-genital shield is always present in the 



32 



THE PLATYSEIINAE (MESOSTI GM AT A : ACEOSEJIDAE) 



male although it may show various degrees of fusion with the ventral shield. The 
genital orifice in this sex is located near the anterior margin of the sterniti-genital. 
The larva has an anal shield usually with a seta on each anal valve in addition to 
the normal three setae present on the shield. There is a tendency for increased 
sckrotization of the anal region in subsequent developmental stages so that the 
protonymph and the deutonymph may have a ventri-anal shield (an anal shield 
with usually one pair of preanals). All the females of the platyseiines we have 
examined have a ventri-anal shield with from one to four pairs of pre-anals. Mem- 
bers of the genus Platyseius in addition have two or more pairs of setae situated 
in the posterior half of the shield (Text-fig. 23). The ventri-anal shield in the 
male shows varying degrees of fusion with the sterniti-genital shield and with the 
dorsal shield (Text-figs. 19 and 31). 




Figs. 12-14. Tarsi of leg I showing development of ambulacra. Fig. 12, Sejus viduus 
C. L. Koch. Fig. 13, Sejus laelaptoides (Berlese). Fig. 14, Sejus unguiculatus (Berlese). 



The larva lacks stigmata and peritremes but the protonymph has a pair of stigmata 
located ventro-laterally between coxae III and IV and short peritremes. In the 
deutonymph, the peritremes are well developed and extend beyond the anterior 
margin of coxae I. The peritrematal shields at this stage, however, are weak and 
inconspicuous. The adults have large peritrematal shields ; these are invariably 
fused with the expodals and extend posterior to coxae IV (Text-fig. 16). Many 
species have a strong poststigmatic prolongation of the peritreme. 

Legs : All the legs are usually long and slender. The structures of taxonomic 
importance are the form of the ambulacrum and the chaetotaxy of tarsi II-IV. 
The ambulacrum of leg I shows varying degrees of development. In a few species 
it is lacking (Text-fig. 12) but generally it comprises a lobate pulvillus and two 
claws. The claws may be minute or relatively large and conspicuous (Text-fig. 13). 
A short pretarsus is usually present ; a notable exception being Sejus unguiculatus 
(Berl.) in which the large claws on tarsus I are sessile (Text-fig. 14). The ambulacra 



THE PLATYSEIINAE (MESOSTIGM AT A : ACEOSEJIDAE) 33 

of legs II-IV are well developed and each consists of a pair of claws and a lobate 
pulvillus. In the majority of the platyseiines, the median and lateral lobes of the 
pulvilli are slender and markedly acute — a probable adaptation for movement over 
a moist substratum. Tarsi II and III and usually IV have a pair of characteristic 
lanceolate setae (Text-fig. 18). Leg II in the male is never crassate or armed with 
spurs. 

CLASSIFICATION 

The characters previously used in separating the genera of the Platyseiinae, e.g. 
the outline of the idiosoma, the presence or absence of an ambulacrum on tarsus I 
and the form of the peritreme, are undoubtedly of some value in the practical 
classification of the subfamily but have no phylogenetic basis. We have concluded 
from our investigations of the external morphology of the group that the chaetotaxy 
and form of the dorsal shield in the adult and immature stages afford the most 
satisfactory characters for a natural classification. It is interesting to note that 
the variation in the form and in the chaetotaxy of the dorsal shield already noted 
for the Aceosejinae (Evans, 1958) is also evident in the Platyseiinae. Thus, there 
is a tendency towards the reduction in the number of the J and S setae on the 
" posterior dorsal shield " and towards the retention of the deutonymphal incisions. 
Our generic concept has as its basis the chaetotaxy of the " posterior dorsal shield " 
which has been discussed above. 



Subfamily Platyseiinae 

Platyseiinae Evans, G. O., (1957), /• Linn. Soc. Lond. (Z00L), 43 : 244. 

Aceosejids with the rostral and internal palptrochanter setae long and usually 
whip-like. Tarsi II and III with a pair of long lanceolate setae about the middle 
of the segment (Text-fig. 18). Dorsal shield in the adults entire, rarely with lateral 
incisions. " Posterior dorsal shield" normally with fifteen pairs of setae; when 
fewer setae are present the J series or S series comprises less than five pairs. Females 
with sternal shield bearing three pairs of setae, a wedge-shaped genital shield with 
or without a pair of setae and a ventri-anal shield with one or more pairs of preanals. 
Male with sterniti-genital and ventri-anal shields, the latter often being partially fused 
with the dorsal shield. Both sexes with or without poststigmatic prolongation of 
the peritreme. Chelicerae chelate-dentate in both sexes ; movable digit with 
spermatophoral process in the male. Tectum basically three pronged. All legs 
usually slender ; leg I with or without ambulacrum, legs II-IV with pulvilli (often 
acutely lobate) and two claws. Leg II unarmed in the male. 



Key to Genera — Adults 

" Posterior dorsal shield " with 5 pairs of setae in the J series (Text-fig. 53) . 
— " Posterior dorsal shield " with only 2 or 3 pairs of setae in the J series (Text-figs. 
15 and 36) .,,,,,,,,,,, 



34 THE PLATYSEIINAE (MESOSTIGMAT A : ACEOSEJIDAE) 

2. " Posterior dorsal shield " with 5 pairs of setae in the S series (Text-fig. 53) ; dorsum 

never with paddle-like setae ; peritreme often with a strong poststigmatic process 
(Text-fig. 54) ; median lobes of the pulvilli of ambulacra II-IV usually acuminate 
(Text-fig. 58) ........ Sejus C. L. Koch (p. 49) 

— " Posterior dorsal shield " with 4 pairs of setae in the S series, ? S2 situated on 
the lateral integument (Text-fig. 179) ; dorsal shield with three or more pairs of 
paddle-like setae, remainder of dorsal setae simple (Text-fig. 184) ; peritreme 
without poststigmatic prolongation ; median lobes of the pulvilli of legs II-IV 
rounded apically (Text-fig. 202) ..... Zerconopsis Hull (p. 91) 

3. Both sexes with three pairs of setae in the J series (Text-fig. 36) ; idiosoma elongate- 

oval in outline ; ventri-anal shield in the female with 3 or 4 pairs of setae in 
addition to the paranal and postanal setae . . . Plesiosejus gen. nov. (p. 42) 

— Both sexes with only 2 pairs of setae in the J series (J4 and J 5) ; idiosoma usually 
subcircular in outline (Text-fig. 15) ; ventri-anal shield in the female with 5 or 6 
pairs of setae in addition to paranal and postanal setae . Platyseius Berlese (p. 34) 



Genus PLATYSEIUS Berlese 

Platyseius Berlese, A., (1916). Redia, 12 : 42. 

Platyseiine mites with the idiosoma usually subcircular in outline. Dorsal shield 
entire : " posterior dorsal shield " with J series comprising two pairs of setae (J4 
and J5). Dorsal setae, with the exception of J5 in some species, long and simple. 
Sternal shield in the female with three pairs of setae ; metasternals situated on 
small plates. Genital shield wedge-shaped and with a pair of setae. Ventri-anal 
shield large and bearing 13 or 15 setae. Poststigmatic process of peritreme and 
peritrematal shield well developed and extending posterior to coxa IV. Male with 
sterniti-genital and ventri-anal shield, and ventri-anal shield showing varying 
degrees of fusion with the dorsal shield. Chaetotaxy of gnathosoma normal. Chelicerae 
chelate-dentate ; movable digit with spermatophoral process in the male. Tectum 
basically trispinate. All legs with ambulacra ; pulvilli of ambulacra II-IV acuminate. 



Type species Lasioseius (Platyseius) capillatus Berlese, 1916 
(=Hypoaspis subglabra Oudemans, 1902) 
Key to Species — Females 

1. Setae i5 shorter than J4 (Text-fig. 15) ; ventri-anal shield with 13 setae ; pre- 

endopodal shields present. Europe . . . h P. subglaber (Oudms.) (p. 35) 

— Setae i5 at least twice the length of J4 (Text-fig. 22) ..... 2 

2. Ventri-anal shield with 15 setae (Text-fig. 27) ; pre-endopodal shields absent. 

Jamaica ........ P. jamaicensis sp. nov. (p. 39) 

— Ventri-anal shield with 13 pairs of setae (Text-fig. 23) ; pre-endopodal shields 

present ............. 3 

3. Setae on ventri-anal shield simple (Text-fig. 23) ; setae J5 thorn-like, smooth. 

Indonesia ......... P. mollicomus Berl. (p. 37) 

— Two pairs of setae on the posterior half of the ventri-anal shield stout, spinose 

(Text-fig. 35) ; setae J 5 short, brush-like. East Africa P. spinosus sp. nov. (p. 41) 






THE PLATYSEIINAE (MESOSTI GM AT A : ACEOSEJIDAE) 

Platyseius subglaber (Oudemans), 1903 



35 



Hypoaspis subglabra Oudemans, A. C, (1903). Ent. Ber., 1 : 87 ; Abh. nat. Ver. Bremen, 18 :88. 
Lasioseius {Platyseius) capillatus Berlese, A., (1916). Redia, 12 : 43, syn. nov. 
Lasioseius (Platyseius) subglaber, Halbert, J. N., (1923). /. Linn. Soc. Lond. (Zool.), 35 : 373. 
Lasioseius (?) listrophorus Schweizer, J., (1924). Arch fur Hydrobiol., 15 : 131. 

Female. Dorsal shield (560 jlc x 361-477 jli) broadly oval ; ornamentation 
restricted to a number of small punctate areas. " Anterior dorsal shield " with 




Figs. 15-18. Platyseius subglaber (Oudemans), female. Fig. 15, dorsum. Fig. 16, venter. 
Fig. 17, chelicera. Fig. 18, tarsus II. 



nineteen pairs of simple setae ; verticals very fine and short (Text-fig. 15). 
" Posterior dorsal shield " with twelve pairs of setae ; J1-J3 being absent. Setae 
J5 short, palmate. The distribution and relative lengths of the setae on the dorsal 
shield and lateral interscutal membrane are shown in the figure. 



36 



THE PLATYSEIINAE (MESO STIGMATA : ACEOSEJIDAE) 



Tritosternum large with a pair of strongly-pilose laciniae, and flanked at its base 
by two pairs of pre-endopodal shields. Sternal shield smooth and with three pairs 
of simple setae (Text-fig. 16). Genital shield wedge shaped and with a pair of 
setae. Ventri-anal shield wider than long (202-215 jlc long x 280-360 /i wide), 
reticulated in its anterior half and bearing thirteen setae. Six small sclerotized 
plates are situated between the genital and ventri-anal shields. Stigma situated 




Figs. 19-21. Platyseius subglaber (Oudemans), male. Fig. 19, venter. 

Fig. 21, tarsus II. 



Fig. 20, chelicera. 



between coxae III and IV ; peritreme extending posterior to the stigma ; peri- 
trematal shield strongly developed. Metapodal shields apparently fused with the 
peritrematal shields. 

Venter of gnathosoma with rostral and internal palptrochanter setae long, 
whip-like. All setae on pedipalp simple. Tectum three-pronged and typical of the 
genus. Dentition of chelicerae as in Text-fig. 17. 



THE PLATYSEIINAE (MESOSTIGM AT A : ACEOSEJIDAE) 37 

Leg I (670-772 pi long) with the tarsus (160-177 l L ) considerably longer than the 
tibia (126-140 fi). Setae on tibia and tarsus fine, those on remaining segments 
stouter. Tarsi II-IV with a pair of long lanceolate setae. Ambulacrum of tarsus 1 
short and claws small. Pulvilli of ambulacra II-IV produced into three long 
acuminate lobes (Text-fig. 18). 

Male. Dorsal shield (425-467 fi x 340-371 /i) is similar to that of the female. 
" Anterior dorsal shield " with nineteen pairs of simple setae and " posterior dorsal 
shield " with twelve pairs of setae. 

The tritosternum is flanked at its base by two or three pairs of small pre-endopodal 
shields. The remainder of the chaetotaxy and sclerotization of the venter is shown 
in Text-fig. 19. 

The gnathosoma and tectum are similar to those of the female. The sperma- 
tophoral process is about three times as long as the movable digit of the chelicera. 
The dentition of the chelicera, and the spermatophoral process are shown in Text- 
fig. 20. 

Leg I (602-682 ft long) has the tarsus (147-177 ju) considerably longer than the 
tibia (114-126 fi). The chaetotaxy of the legs is as in the female. The ambulacra 
of tarsi II-IV are provided with lanceolate pulvilli (Text-fig. 21). 

Distribution. This species is widely distributed in western Europe. In Great 
Britain we have examined specimens from Cumber Park, Nottinghamshire (coll. 
G. O. E., April, 1955), Dungeness, Kent (coll. D. A. Chant, July, 1955), Box Hill, 
Surrey (coll. E. Duffey, 30.vi.1951), and St. Agnes, Isles of Scilly (coll. K. H. H., 
September, 1957). 



Platyseius mollicomus Berlese, 1916 

Lasioseius {Platyseius) mollicomus Berlese, A., (1916). Redia, 12 : 43. 

Platyseius mollicomus, Vitzthum, H. C, (1931). Arch, fur Hydrobiol., Suppl., 9 : 70, figs. 

Female. Dorsal shield (570-580 ju x 467-480 jli) broadly oval in outline ; 
ornamentation restricted to a number of faint reticulated areas. " Anterior dorsal 
shield " with twenty-one pairs of simple setae (Text-fig. 22). " Posterior dorsal 
shield" with twelve pairs of simple setae; J1-J3 being absent. Setae J5 are 
short and simple. The distribution and relative lengths of the setae on the dorsal 
shield are shown in the Text-fig. 

Tritosternum large with a pair of strongly pilose laciniae, and flanked at its base 
by two pairs of pre-endopodal shields. Sternal shield with three pairs of simple 
setae and faintly reticulated (Text-fig. 23). Genital shield slightly convex pos- 
teriorly and with a pair of setae. Ventri-anal shield wider than long (177 /i long 
X 318 jlc wide) with a transverse line anteriorly, and bearing thirteen setae. Four 
small sclerotized plates are situated between the genital and ventri-anal shields. 
Stigma situated between coxae III and IV. Peritreme broad, occupying most of 
the peritrematal shield, and with two deep indentations (Text-fig. 24) , and extending 
posterior to the stigma. Metapodal shields fused with the peritrematal shields. 

Venter of gnathosoma with rostral and internal palptrochanter setae long and 



38 



THE PLATYSEIINAE (MESOSTI GM AT A : ACEOSEJIDAE) 



whip-like. All setae on pedipalp simple. Tectum three-pronged and typical of 
the genus. The chelicerae are typical. 

Leg I (700-760 ju, long) with the tarsus (202 fi) longer than the tibia (126 /j). Setae 
on tarsus fine, those on remaining segments stouter. Tarsi II-IV with a pair of 
long lanceolate setae. Ambulacrum of tarsus I fairly short and claws small. 
Pulvilli of ambulacra II-IV produced into three acuminate lobes (Text-fig. 25). 




Figs. 22-25. Platyseius mollicomus Berlese, female. Fig. 22, dorsum. Fig. 23, venter. 
Fig. 24, peritrematal shield. Fig. 25, tarsus II. 



Male. This sex (415 /i x 340 /j) is described by Vitzthum (1931) who also gives 
a figure of the chelicera. 

Distribution. Berlese (1916) based his description on material collected in 
Java by CI. Jacobson. Vitzthum (1931) examined specimens (male, female and 
deutonymph) from Tjibodas, Java (Indonesia). The Museum Collections contain a 
female and a deutonymph collected by A. H. G. Alston in decaying leaves and 



THE PLATYSEIINAE (MESOSTIGM AT A : ACEOSEJIDAK) 



39 



fruit, Bogor, Indonesia (1956.2.27.6) ; and a single female collected by G. Owen 
Evans in grassland nine miles north-east of Bundibugyo, Ruwenzori, Uganda, 
24.viii.1952 (1959. 1.20. 1). 



Platyseius jamaicensis sp. nov. 

Female. Dorsal shield (498 jjl x 392-402 ju,) broadly oval in outline ; ornamenta- 
tion restricted to a number of small punctate areas. "Anterior dorsal shield" 
with twenty pairs of simple setae of which the verticals are the shortest (Text-fig. 




Figs. 26-30 Platyseius jamaicensis sp. nov., female. Fig. 26, dorsum. Fig. 27, venter. 
Fig. 28, tectum. Fig. 29, chelicera. Fig. 30, tarsus II. 



26). " Posterior dorsal shield " with thirteen pairs of setae ; J1-J3 being absent. 
Setae J5 short, palmate and inconspicuous. J4 and Z5 are stout. The distribution 
and relative lengths of the setae on the dorsal shield are shown in the Text-fig. 

Tritosternum large with a pair of strongly-pilose laciniae. Sternal shield with a 
pair of longitudinal elongate punctate areas and three pairs of simple setae (Text- 
fig. 27). Genital shield convex posteriorly and finely reticulated, with a pair of 
setae. Ventri-anal shield wider than long (152 /i long x 253-273 /i wide), reticulated 



4 o 



THE PLATYSEIINAE (MESOST IGM AT A : ACEOSEJIDAE) 



anteriorly and bearing fifteen setae. Four small sclerotized plates are situated 
between the genital and ventri-anal shields. Stigma situated between coxae III 
and IV, peritreme extending posterior to the stigma ; peritrematal shield strongly 
developed. Metapodal shields apparently fused with the peritrematal shields. 

Venter of gnathosoma with rostral and internal palptrochanter setae long, 
whip-like. All setae on pedipalp simple. Tectum (Text-fig. 28) three-pronged 




Figs. 31-33. Platyseius jamaicensis sp. nov., male. Fig. 31, venter. Fig. 32, tectum. 

Fig. 33, chelicera. 



and typical of the genus. Dentition of chelicera as in Text-fig. 29. 

Leg I (about 770 /i long) with the tarsus (177 /i) longer than the tibia (157 fi). 
Setae on tibia and tarsus very fine : those on remaining segments stouter. Tarsi 
II-IV with a pair of long lanceolate setae. Ambulacrum of tarsus I short and the 
claws small. Pulvilli of ambulacra II-IV produced into three acuminate lobes 
(Text-fig. 30). 



THE PLATYSEIINAE (MESOSTI GM AT A : ACEOSEJIDAE) 



4i 



Male. Dorsal shield (498 /i X 310 ju) similar to that of the female. The tri- 
tosternum is of the same form as in the female. The chaetotaxy and sclerotization 
of the venter are shown in Text -fig. 31. 

The gnathosoma is similar to that of the female, and the tectum is shown in 
Text-fig. 32. The spermatophoral process is about twice as long as the movable 
digit of the chelicera (Text-fig. 33). 

Leg I (c. 680 pi) with the tarsus and tibia each 152 fi. The chaetotaxy and 
ambulacra of the legs are as in the female. 

Locality. Jamaica, B.W.I. The holotype female (1959. 1.20. 18) and one 
paratype from Fairy Glade, St. Andrew, in Lycopodium, 4,300 ft., 6.xii.i956; 
and the allotype male (1959. 1.20. 19) from St. Thomas, John Cove Mountains, in 
moss, 2,300 ft., 25. v. 1956 collected by P. F. Bellinger. The paratype female has 
been returned to the collector. 



Platyseius spinosus sp. nov. 

Female. Dorsal shield (498 pi x 402 pi) broadly oval in outline ; ornamentation 
restricted to a number of very faint reticulated areas. " Anterior dorsal shield " 




Figs. 34-35. Platyseius spinosus sp. nov., female. Fig. 34, dorsal shield. Fig. 35, venter. 



with twenty-one pairs of simple setae (Text-fig. 34). Posterior dorsal shield with 
thirteen pairs of simple setae ; J1-J3 being absent. Setae J5 are short, palmate and 
inconspicuous. Z5 are stout. The distribution and relative lengths of the setae 
on the dorsal shield are shown in the Text-fig. 

ZOOL. 6, NO. 2. 



\ 



42 THE PLATYSEIINAE (MESOSTIGMAT A : ACEOSEJIDAE) 

Tritosternum large with a pair of strongly-pilose laciniae, and flanked at its base 
by one or two pairs' of pre-endopodal shields. Sternal shield with three pairs of 
simple setae and with a characteristically-shaped central mark (Text-fig. 35). 
Genital shield slightly convex posteriorly and with a pair of setae. Ventri-anal 
shield wider than long (177 ju, long x 298 jlc wide) with a transverse line anteriorly 
and bearing thirteen setae of which two pairs are stout. A narrow sclerotized 
band and two narrow plates are situated between the genital and ventri-anal shields. 
Stigma situated between coxae III and IV, peritreme broad, occupying most of 
the peritrematal shield, with a deep indentation, and extending posterior to the 
stigma. Metapodal shields fused with the peritrematal shields. 

Venter of gnathosoma with rostral and internal palptrochanter setae long, whip- 
like. All setae on pedipalp simple. Tectum three-pronged and typical of the 
genus. The chelicerae are typical. 

Leg I (707 jll long) with the tarsus (202 ju) longer than the tibia (126 /i). Setae 
on tarsus fine ; those on remaining segments stouter. Tarsi II-IV with a pair of 
long lanceolate setae. Ambulacrum of tarsus I short and claws small. Pulvilli 
of ambulacra II-IV produced into three acuminate lobes. 

Male. Unknown. 

Locality. Ruwenzori, Uganda. A single female (1959. 1.20.20), collected by 
G. Owen Evans, in grassland at 2,850 ft., 24.viii.1952, nine miles north-east of 
Bundibugyo. 



Genus PLESIOSEJUS gen. nov. 

Platyseiine mites with the " posterior dorsal shield " having only three pairs of 
setae in the J series (Ji and J2 absent). Z and J series each with five pairs. Lateral 
margins of the dorsal shield entire in the adult stages. Vertical setae present or 
absent. Sternal shield in the female with three pairs of simple setae ; metasternals 
free. Genital shield wedge-shaped and bearing a pair of setae. Ventri-anal shield 
large with three or four pairs of setae in addition to the paranals and the postanal 
seta. Male with sterniti-genital and ventri-anal shields, the latter showing varying 
degrees of fusion with the podal-peritrematal and dorsal shields. 



Type species Ameroseius italicus Berlese, 1905 
Key to Species — Females 

Without poststigmatic prolongation of peritreme (Text-fig. 50) ; majority of dorsal 

setae long, stout (Text-fig. 49) ; vertical setae well developed, directed anteriorly ; 

ventri-anal shield with 9 setae. Jamaica . . P. horridus sp. nov. (p. 48) 

— With strong poststigmatic prolongation of peritreme extending posterior to coxa 

IV ; dorsal setae considerably shorter (Text-fig. 36) ; vertical setae absent ; 

ventri-anal shield with 1 1 setae ......... 2 

Dorsal shield with distinct " tubercle " posterior to J4 (Text-fig. 36). Europe 

P. italicus (Berlese), (p. 43) 
— Dorsal shield without such " tubercle " (Text-fig. 42). P. major (Halbert), (p. 47) 



THE PLATYSEIINAE (MESOSTIGM ATA : ACEOSEJIDAE) 43 



Plesiosejus italicus (Berlese), comb. nov. 

Ameroseius italicus Berlese, A., (1905). Redia, 2 : 234 ; (1906). Ibid., 6 : tav. 19, Text-figs. 

35, 35a. 
Paraseius tenuipes Halbert, J. N. (1915). Proc. R. Irish Acad., 39ii : 78, syn. nov. 
Lasioseius italicus Berlese, A., (1916). Redia, 12 : 34. 

t Lasioseius (Episeius) michaeli Halbert, J. N., (1923). /. Linn. Soc. Lond. (Zool.), 35 : 373. 
Episeius tenuipes, Schweizer, J., (1949). Res. Rech. sci. Pare Nat. Suisse, N.F. 2 : 60. 
E. tenuipes, Franz, H., (1943). Denkschr. Akad. Wiss. Wien., 107 : 87. 

Female. Dorsal shield (550-651 jli long x 310-402 jli wide) oval; ornamentation 
comprising a reticulated pattern covering the whole shield (Text-fig. 36). " Anterior 
dorsal shield " with twenty pairs of simple setae ; verticals absent. " Posterior 
dorsal shield" with thirteen pairs of setae; J1-J2 being absent; J5 are finely 
pilose. A strong tubercle arising posterior to setae J3 denotes the highest point 
on the median ridge of the dorsal shield. The distribution and relative lengths of 
the setae and the structure of the dorsal shield are shown in the Text-fig. 

Tritosternum well developed with pilose laciniae, and flanked at its base by one 
or two pairs of small pre-endopodal shields. Sternal shield with three pairs of 
simple setae and two oval areas antero-medially (Text-fig. 37). Genital shield 
convex posteriorly, and bearing one pair of setae. Ventri-anal shield considerably 
wider than long (177-202 jli long X 265-341 jlc wide), reticulated and bearing eleven 
simple setae. Between the genital and ventri-anal shields lie six narrow sclerotized 
plates. Stigma between coxae III and IV and the peritreme extending posterior to 
coxa IV ; peritrematal shield strongly sclerotized. Metapodal shields free and 
transversely situated. 

Venter of gnathosoma with rostral and internal palptrochanter setae long, whip- 
like. All setae on pedipalp simple. Tectum three-pronged and typical of the 
genus. The dentition of the chelicera is shown in Text-fig. 38. 

Leg I (639-726 jli long) with the tarsus (167-195 jli) longer than the tibia (113- 
129 jli). Setae on tarsus and tibia very fine ; those on remaining segments stouter ; 
ambulacra short and claws small. Tarsi II-IV with a pair of lanceolate setae ; 
pulvilli produced into three acuminate lobes (Text-fig. 39). 

Male. Dorsal shield (467-507 jli long x 279-310 jli wide) entirely reticulated as 
in the female. "Anterior dorsal shield" with nineteen pairs of simple setae, 
verticals absent. " Posterior dorsal shield" with thirteen pairs of setae; J1-J3 
absent ; J5 short and pilose. Postero-dorsal tubercle not so pronounced as in the 
female. 

Tritosternum as in the female. The chaetotaxy and sclerotization of the venter 
are shown in Text-fig. 40. The gnathosoma and tectum are similar to those of the 
female. Spermatophoral process four times as long as the movable digit of the 
chelicera (Text-fig. 41). 

Leg I (534 jli long) with the tarsus (147-152 /i) longer than the tibia (96-106 /i). 
The chaetotaxy and ambulacra of the legs are similar to the female. 



44 



THE PLATYSEIINAE (MESOSTIGMATA : ACEOSEJIDAE; 



Distribution. Berlese's (1905 and 1916) descriptions of the male and female of 
italicus are based on specimens collected under dead rotting leaves on a marsh at 
Udine in northern Italy. Halbert (191 5) described P. tenuipes from a single female 
" found amongst moss on a stone in a mountain stream at Glencree, County Dublin, 
May ". Schweizer (1949) records tenuipes from the Swiss National Park. We have 




Figs. 36-39. Plesiosejus italicus (Berlese), female. Fig. 36, dorsum. Fig. 37, venter. 
Fig. 38. chelicera. Fig. 39, tarsus II 



THE PLATYSEIINAE (MESOSTI GM AT A : ACEOSEJIDAE) 



45 



examined British specimens from the following localities : Thirlmere, Cumberland, 
25. ix. 1956, coll. P. N. Lawrence ; Grasmere, Westmorland, 29.vi.1951, coll. J. T. 
Salmon; Blelham Tarn, Lancashire, 22. hi. 1955, coll. R. J. Elliott; summit of 
Snowdon, Caernarvonshire, August, 195 1, coll. E. Duffey ; Minworth, Birmingham, 
10. ix. 1952, coll. T. G. Tomlinson ; Swithland Wood and Anstey, Leicestershire, 
19. iv. 1958, coll. P. N. L.; Sea Houses, Northumberland, i.x.1951, coll. J. T. 





Figs. 40-41. Plesiosejus italicus (Berlese), male. Fig. 40, venter. Fig. 41, chelicera. 



Salmon; Oxted, Surrey, 20. i. 1957, coll. P. N. L.; Leith Hill, Surrey, 6.VL1951, 
coll. J. T. Salmon; Keston, Kent, 2.viii.i957, coll. P. N. L. ; Dungeness, Kent, 
8.vii.i955, coll. D. A. Chant; Avonmouth, Gloucestershire, 11. v. 1957, coll. 
P. N. L.; Combe Martin, Devonshire, March and June, 1957, coll. P. N. L. and 
M. E. Bacchus ; Porth Gwarra, Cornwall (A. D. Michael Coll.) ; The Burren, Co. 
Clare, 6.VL1951, coll. E. A. J. Duffy. 



46 THE PLATYSEIINAE (MESOSTIGM ATA : ACEOSEJIDAE; 




Figs. 42-47. Plesiosejus major (Halbert). Fig. 42, dorsum of female. Fig. 43, venter 
of female. Fig. 44, chelicera of female. Fig. 45, tarsus II of female. Fig. 46, venter 
of male. Fig. 47, chelicera of male. 



THE PLATYSEIINAE (MESOSTIGM AT A : ACEOSEJIDAE) 47 



Plesiosejus major (Halbert), comb. nov. 

Lasioseius (Episeius) major Halb.ert, J. N., (1923). /. Linn. Soc. Zool. Lond. 35 : 373. 
Lasioseius (L.) tenuipes, Schweizer, J., (1922). Verh. Naturf. Ges. Basel, 33 : 43. 
Episeius major, Schweizer, J., (1949). Res. Rech. sci. Pare Nat. Suisse, N.F. 2 : 61. 

Female. Dorsal shield (581-740 jlc long x 371-488 jn wide) broadly oval in 
outline : ornamentation consisting of a dense reticulated pattern covering the whole 
of the shield (Text-fig. 42). " Anterior dorsal shield " with twenty pairs of simple 
setae. " Posterior dorsal shield " with thirteen pairs of setae ; J1-J2 being absent ; 
setae J5 are short and spiculate. The distribution and relative lengths of the dorsal 
setae are shown in the Text-fig. 

Tritosternum well developed with pilose laciniae, and flanked at its base by two 
or three pairs of small pre-endopodal shields. Sternal shield with three pairs of 
simple setae (Text-fig. 43) . Genital shield with one pair of setae and slightly convex 
posteriorly. Ventri-anal shield wider than long (203-253 /i long X 288-379 ju, wide), 
strongly reticulated and bearing eleven simple setae. Six narrow sclerotized plates 
are situated transversely on the membrane between the genital and ventri-anal 
shields. Stigma situated between coxae III and IV with the peritreme extending 
posterior to coxa IV. Peritrematal shield strongly sclerotized. Metapodal shields 
free and obliquely situated. 

Venter of gnathosoma with rostral and internal palptrochanter setae long, whip- 
like. All setae on pedipalp simple. Tectum three-pronged and typical of the genus. 
The dentition of the chelicera is shown in Text-fig. 44. 

Leg I (681-797 fi long) with the tarsus (147-177 /i) longer than the tibia (106- 
152 fi) ; pulvillus short, claws small. Setae on tarsus and tibia very fine ; those 
on remaining segments stouter. Tarsi II-IV with a pair of lanceolate setae ; 
pulvilli produced into three acuminate lobes (Text-fig. 45). 

Male. Dorsal shield (550 fi long x 340 /i wide) of the same form and chaetotaxy 
as the female. The structure and chaetotaxy of the venter are shown in Text-fig. 
46. The gnathosoma and tectum are similar to those of the female. The sperma- 
tophoral process is twice as long as the movable digit of the chelicera (Text-fig. 47). 

Leg I (670 jlc long) with the tarsus (165 ju) longer than the tibia (126 /i). The legs 
are similar to those of the female. 

Distribution. Halbert's (1923) description of males and females of this species 
is based on specimens found in moss in streams at Kilmashogue and Glencullen in 
Ireland, and at Mill Bay, Land's End, England. Schweizer (1922) described what 
he considered to be males of Lasioseius tenuipes (Halbert) from Basel, Switzerland, 
but his figures show clearly that his specimens are Plesiosejus major (Halbert). In 
1949 the same author recorded major from the Swiss National Park. We have 
examined specimens collected by the River Tolka, Glasnevin, Co. Dublin (Halbert 
Collection), and from the following localities: Oxted, Surrey, 20. i. 1957, coll. 
P. N. Lawrence ; Cumber Park, Nottinghamshire, April, 1955, coll. G. O. E. ; 
Markfield and Anstey, Leicestershire, 19. iv. 1958, coll. P. N. L.; Blea Tarn, West- 



4 8 



THE PLATYSEIINAE (MESOSTI GM AT A : ACEOSEJIDAE) 



morland, 27. ix. 1956, coll. P. N. L.; and Chorley, Lancashire, 21. xi. 1940, coll. 
Water Pollution Research Laboratory. All these British specimens were collected 
from wet habitats (mosses, etc.). 



Plesiosejus horridus sp. nov. 

Female. Dorsal shield (350 /i long x 218 /i wide) narrower in outline than in 
the other species of the genus, and is steeply elevated posteriorly (Text-fig. 48). 




Figs. 48-52. Plesiosejus horridus sp. nov., female. Fig. 48, lateral view of dorsum. 
Fig. 49, dorsal shield. Fig. 50, venter. Fig. 51, chelicera. Fig. 52, tarsus II. 



There is no noticeable ornamentation on the dorsal shield. " Anterior dorsal 
shield " with twenty pairs of setae. J1-J3 absent. Setae J5 are partly concealed 
by the postero-dorsal elevation of the dorsal shield which bears J4 at its apex 
(Text-fig. 49). The distribution and relative lengths of the dorsal setae are shown 
in the Text-fig. 

Tritosternum well developed with pilose laciniae. Sternal shield with three pairs 



THE PLATYSEIINAE (MESOSTIGM AT A : ACEOSEJIDAE) 49 

of short simple setae (Text-fig. 50) . Genital shield flask shaped with a pair of setae. 
Ventri-anal shield slightly wider than long (93 fi long x 113 fi wide), with nine setae 
of which two lateral pairs are long. There appears to be a narrow sclerotized band 
between the genital and ventri-anal shields. Stigma between coxae III and IV, 
with the narrow, crenate peritreme not extending posterior to the stigma. Peri- 
trematal shield extending to the posterior margin of coxa IV. Met apodal plates 
oval, free. 

Venter of gnathosoma with rostral and internal palptrochanter setae long, whip- 
like. All setae on pedipalp simple. Tectum three-pronged and typical of the 
genus. The dentition of the chelicera is shown in Text-fig. 51. 

Leg I (370 /i long) with the tarsus (96 /jl) one-and-a-half times as long as the tibia 
(63/j). Setae on tarsus very fine ; those on remaining segments stouter. Tarsi II- 
IV with a pair of lanceolate setae. Tarsus I without claws. Pulvilli of tarsi II-IV 
produced into three acuminate lobes (Text-fig. 52). Leg IV 370 fi long with the 
tarsus 130 fi. 

Male. Unknown. 

Locality. Jamaica, B.W.I. The holotype female (1959. 1.20. 41) and two 
paratype females (1959. 1.20. 42) in damp leaf litter, St. Ann, Mt. Diablo, 2,200 ft., 
3.VL1956, collected by P. F. Bellinger. One paratype has been returned to the 
collector. 



Genus SEJUS C. L. Koch 

Sejus Koch, C. L., (1843) 1 Ubers. d. Arachnidensyst., Heft 3 : 92. 
Cheiroseius Berlese, A., (1916). Redia, 12 : 33. 

Episeius Hull, J. E., (1918). Trans, nat. Hist. Soc. Northumb. (n.s.), 5 : 63. 
Episeiella Willmann, C, (1938). Ann. Hist. nat. Mus. Hung., 31 : 167. 

Platyseiine mites with the " posterior dorsal shield " bearing fifteen pairs of setae 
of which five pairs belong to the J series ; S series also comprising five pairs. Dorsal 
setae usually simple, rarely pilose. Lateral margins of the dorsal shield entire or 
incised. Sternal shield in the female with three pairs of setae ; metasternals free. 
Genital shield wedge shaped and bearing a pair of setae. Ventri-anal shield large 
with two or more pairs of pre-anals. Peritreme with or without poststigmatic 
process. Leg I rarely without ambulacrum, usually with at least two claws. Legs 
II-IV with normal ambulacrum. 

This is, at present, the largest genus of the subfamily. A number of the older 
species are inadequately described and must remain species incertae sedis until the 
type material can be re-examined. The following key includes only those species 
we have examined. 



Type species Sejus viduus C. L. Koch, 1839 

1 PP- 73 _I 3 2 °f the third Heft were published in 1843 although most authors consider the complete 
Heft to have appeared in 1842. 



5 o THE PLATYSEIINAE (MESOSTIGM AT A : ACEOSEJIDAE) 

Key to the Species in the British Museum (Natural History) — Females 

i. Dorsal shield with lateral incisions (Text-figs. 114 and 1 68) .... 2 

— Lateral margins of the dorsal shield entire ....... 3 

2. Lateral incisions located anterior to setae Si and coinciding with their position in 

the deutonymph (Text-fig. 114) ; median process of tectum long, Y-shaped 
(Text-fig. 117) ; vertical setae absent; ventri-anal shield with one pair of 
preanals (Text-fig. 116). Sikkim .... 5. clayi sp. nov. (p. 73) 

—Lateral incisions wide and situated between S2 and S3 (Text. -fig 168) ; median 
process of tectum not Y-shaped ; vertical setae present ; ventrianal shield 
with two pairs of preanals (Text-fig. 169). Argentina . 5. ornatus sp. nov. (p. 88) 

3. Tarsus I without ambulacrum (Text-fig. 12) . . . . . . . 4 

— Tarsus I with ambulacrum or at least with two distinct claws ... 6 

4. Vertical setae considerably longer than i2 (Text-fig. 53) ; poststigmatic process of 

the peritreme extending posterior to coxa IV (Text-fig. 54). Europe 

5. viduus C. L. Koch. (p. 52) 
— Vertical setae shorter than i2 (Text-fig. 162) ; poststigmatic process very short 

or absent ............. 5 

5. With a short but conspicuous poststimatic process of the peritreme ; sternal shield 

with an elongate reticulated area (Text-fig. 164). Sikkim and Argentina 

5. browningi sp. nov. (p. 86) 
— Without a poststigmatic process ; sternal shield without such ornamentation 

(Text-fig. 144). Jamaica ...... S. hulli sp. nov. (p. 81) 

6. Peritreme not produced posterior to the stigma (Text-fig. 81) or poststigmatic 

process narrow, short, never extending posterior to coxa IV (Text-fig. 154) . 7 

— Poststigmatic portion of the peritreme strongly developed and extending posterior 

to coxa IV (Text-fig. 56) ......... 17 

7. Dorsal shield strongly arched (Text-fig. 79) ; sternal region not definitely sclero- 

tized (Text-fig. 81) ; a number of dorsal setae broad, pilose ; ventrianal shield 
with 7 setae. Britain ..... 5. dromadis sp. nov. (p. 62) 

— Dorsal shield at the most with a median longitudinal ridge ; sternal shield well- 

sclerotized ; dorsal setae simple or lanceolate ...... 8 

8. Leg IV greatly elongated, more than twice the length of the dorsal shield (Text-fig. 

99) ; vertical setae prominent, directed anteriorly ; ventri-anal shield with 9 
setae. E. Africa ...... S. phalangioides sp. nov. 

— Leg IV never more than i| times the length of the dorsal shield 

9. Vertical setae apparently absent (Text -fig. 124) ; dorsal and ventral shields charac- 

teristically ornamented (Text-figs. 124, 125, 129). Jamaica 

S. aciculatus sp. nov. 
— Vertical setae present, prominent, directed anteriorly (Text-fig. 153) 

10. Peritreme with a distinct poststigmatic process (Text-fig. 154) .... 
— Peritreme without a poststigmatic process (Text-fig. 141) 

11. Dorsal shield with a distinct " tubercle " in the region of setae J 4 — this indicates 

the highest point of the median longitudinal ridge (Text-fig. 153) ; postigmatic 
portion of peritreme extending to about the middle of coxa IV. Jamaica 

5. neborealis sp. nov. 
— Dorsal shield without tubercle in the region of setae J4 . 

12. Vertical setae shorter than 32 (Text-fig. 103) ; femur, genu and tibia of leg with 

sharply pointed spine-like setae situated on tubercles along their anterior and 
posterior margins ; ventri-anal shield with 9 setae (Text-fig. 104). E. Africa 

S. spinipes sp. nov. (p. 69) 
— Vertical setae long, about equal in length to J2 (Text-fig. 63) ; chaetotaxy of 

leg I normal ; ventri-anal shield with 11 setae. Europe S. borealis (Berl.) (p. 56) 



(p. 66) 
9 


(P- 76) 
10 


11 


13 


(p. 84) 
12 



15 


(P- 85) 


(P 81). 


(P. 65) 



THE PLATYSEIINAE (MESOSTIGM AT A : ACEOSEJIDAE) 51 

13. Tarsus I with a long, strong seta situated dorsally towards the middle of the 

segment (Text-fig. 140) . . . . . . 

— Tarsus I without such a prominent seta dorsally ...... 

14. Dorsal shield with a conspicuous tubercle overhanging setae J5 (Text-fig. 158). 

Jamaica ...:.... 5. nodosus sp. nov. 

— Dorsal shield without a tubercle in the region of setae J4 and J5 (Text-fig. 138). 
Jamaica ........ S. bellingeri sp. nov. 

15. Ventri-anal shield broad, gently rounded posteriorly (Text-fig. 92) ; peritrematal 

shield not developed posterior to coxa IV. E. Africa . 5. kennedyi sp. nov. 
— Ventri-anal shield markedly tapering in its posterior half (Text-figs. 121 and 

132), subtriangular in outline ......... 16 

16. Sternal shield with a distinct reticulated area medially ; pre-endopodal shields 

absent ; paranal setae situated on a strongly-sclerotized bar (Text-fig. 132). 
Jamaica ........ 5. antillanus sp. nov. (p. 78) 

— Sternal shield without reticulated area ; pre-endopodal shields present (Text- 
fig. 121) ; paranals not on sclerotized bar. Indonesia . S. alstoni sp. nov. (p. 74) 

17. Claws on leg I large, sessile (Text-fig. 14) ; tarsus I with a thick spine on its 

anterior margin. Europe .... 5. unguiculatus (Berl.) (p. 53) 

— Claw on leg I normal, usually situated on a pretarsus, if sessile claws minute 

(Text-fig. 13) 18 

18. Surface of dorsal shield minutely punctured or tuberculated (Text-fig. 55), reticula- 

tion when present restricted to the lateral margins of the shield . . . 19 
— Surface of the dorsal shield reticulated, often scabrid, without punctures (Text- 
fig- 74) 2 ° 

19. Dorsal shield minutely punctured (Text-fig. 55) ; sternal shield with an oval 

reticulated area (Text-fig. 56) ; ventri-anal shield with 11 setae. Europe 

5. laelaptoides (Berl.) (p. 54) 
— Dorsal shield tuberculated (Text-fig. 174) ; sternal shield lacking a reticulated 
area ; ventri-anal shield with 9 setae (Text-fig. 175). Argentina 

5. tuberculatus sp. nov. (p. 89) 

20. Claws on leg I minute, apparently sessile ; leg I long and slender, about 1% times the 

length of the dorsal shield .......... 21 

Claws on leg I situated on a short but distinct pretarsus ; leg I usually considerably 

less than ii times the length of the dorsal shield . . . . . . 22 

2 1 . Tibia and tarsus I approximately equal in length ; tarsus I more than 200// in length. 

Jamaica ........ S. jamaicensis sp. nov. (p. 83) 

— Tibia I distinctly longer than tarsus I ; tarsus I not more than 160 p in length. 

Nepal ........ 5. nepalensis sp. nov. (p. 71) 

12. Vertical setae long, directed anteriorly (Text-fig. 74) ; dorsal shield heavily orna- 
mented ; tarsus I about twice the length of tibia I. British Isles 

5. cassiteridum sp. nov. (p. 61) 
— Vertical setae relatively shorter, directed antero-laterally or posteriorly (Text- 
figs. 67 and 86) ; vertex not well developed ....... 23 

»3. Tibia I equal in length to or longer than tarsus I. Europe S. serratus (Halbert) (p. 58) 
Tibia I distinctly shorter than tarsus I (1:1.3-1.7). . . . . . 24 

'4. Ventri-anal shield with five subcircular areas arranged crescentically posterior to 
the paranals (Text-fig. 87) ; small weakly-sclerotized species about 360// in 
length. E. Africa ....... 5. areolatus sp. nov. (p. 64) 

Ventri-anal shield without such areas ; larger species 400-800^ in length . . 25 

-5- J 5 about I the length of J4, setiform (Text- fig. no). Nepal 

5. parbatensis sp. nov. (p. 72) 
J 5 less than £ the length of J 4, usually thorn-like . . . . . . 26 



52 



THE PLATYSEIINAE (ME SOSTI GM AT A : ACEOSEJIDAE) 



26. Sternal shield without semicircular depressions near its anterior margin (Text-fig. 

71). Europe ........ 5. curtipes (Halbert) (p. 59) 

— Sternal shield with a pair of semicircular depressions near its anterior margin 

(Text-fig. 60) ........... . 27 

27. Sclerotized bases of the vertical setae large, almost contiguous (Text-fig. 59) ; 

highest point of median dorsal ridge located between i5. Europe 

5. necorniger (Oudemans) (p. 55) 
— Bases of vertical setae weakly sclerotized, widely separated (Text-fig. 100) ; 
highest point of median dorsal elevation located between 15 and Ji. E. Africa 

S. signatus sp. nov. (p. 68) 



Sejus viduus C. L. Koch 

Sejus viduus Koch, C. L., 1839, Deutsch. Crust. Myr. Arach., Fasc. 24, t. 10. 

Epicrius comiger Berlese A., 1891, A. M.S., Fasc. 59, No. 2. 

Sejus viduus C. L. Koch, Oudemans, A. C, 1936, K.H.O.A., 3A : 376. 

Episeiella heteropoda Willmann, C, 1938, Ann. Hist. nat. Mus. Hung., 31 : 164, syn. nov. 

Female. Dorsal shield (529 /i long x 330 fi wide) covered by a strong irregular 




53' 

Figs. 53-54. Sejus viduus (C. L. Koch), female. Fig. 53, dorsal shield. Fig. 54, venter. 






THE PLATYSEIINAE (MESOSTIGM AT A : ACEOSEJIDAE) 53 

network. Lateral margins irregular. "Anterior dorsal shield" with twenty-one 
pairs of simple setae ; verticals very prominent, well separated, and directed 
anteriorly. " Posterior dorsal shield " with fifteen pairs of simple setae of which 
J5 are the shortest. All the setae arise from small protuberances (Text-fig. 53). 

Tritosternum with a pair of pilose laciniae. Sternal shield with a reticulated area 
medially and bearing three pairs of simple setae. Genital shield slightly convex 
posteriorly and bearing a single pair of simple setae. Ventri-anal shield broader 
than long (101 fi long X 139 fi wide) with only faint traces of reticulations, and 
bearing nine simple setae. Between the genital and ventri-anal shields lie four 
narrow sclerotized platelets. Stigma situated between coxae III and IV ; posterior 
prolongation of peritreme, and peritrematal shield, reaching beyond the posterior 
margin of coxa IV (Text-fig. 54). Metapodal shields small. 

Venter of gnathosoma with rostral and internal palptrochanter setae long, whip- 
like. All setae on pedipalp simple. Tectum three-pronged, and chelicerae typical 
of the genus. 

Leg I (724 [i long) with the tarsus (144 /i) shorter than the tibia (167 fi). Setae 
on tarsus, tibia and genu I very fine ; tarsal claws absent ; remaining setae stouter. 
Tarsi II-IV with a pair of lanceolate setae ; ambulacra without acuminate lobes. 

Male. Unknown. 

Distribution. Koch's (1839) description is based on material from wet woodland 
habitats in Germany. Berlese's (1891) description of Epicrius corniger is based on 
several specimens in moss on high mountains, Venice. Berlese's type has been 
examined by us in Florence. Willmann's (1939) description of Episeiella heteropoda, 
was made from one female sifted at Vihnyei volgy, Hungary, and is the specimen 
from which our description and figures are made. 



Sejus unguiculatus Berlese 

Sejus unguiculatus Berlese, A., 1887, A. M.S., 41, No. 4. 

Lasioseius (Cheiroseius) unguiculatus Berlese, A., 1916, Redia, 12 : 33. 

The characteristic feature of this species is that the claws on tarsus I are greatly 
enlarged and sessile (Text-fig. 14). The chaetotaxy of the dorsal shield is typical of 
the genus. The " posterior dorsal shield " bears fifteen pairs of setae ; five pairs 
in the J series. The " anterior dorsal shield " bears twenty pairs of setae ; vertex 
weakly developed. The chaetotaxy and sclerotization of the venter are typical of 
Sejus. Sternal shield with three pairs of setae ; genital shield with one pair of 
setae ; ventri-anal shield large, occupying most of the area posterior to coxae IV, 
and bearing eleven simple setae. Six small plates are situated between the genital 
and ventri-anal shields. Peritreme extending posterior to coxa IV. Metapodal 
plates conspicuous. 

Berlese described this species from moss in Venice. The above description is 
based on drawings of the type. This species is not represented in the Museum 
Collections. 



54 



THE PLATYSEIINAE (MESOSTIGMATA : ACEOSEJIDAE) 



Sejus laelaptoides (Berlese) comb. nov. 

Epicrius laelaptoides Berlese, A., 1887, A. M.S., Fasc. 40, No. 10. 

Parasejus glaber var. minor Tragardh, I., 1910, Naturw. Untersuch. Sarekgeb., 4 : 432, syn. nov. 

Lasioseius (Episeius) sphagni Halbert, J. N., 1923, /. Linn. Soc. Lond. (Zool.), 35 : 371, syn. nov. 

Female. Dorsal shield (518-581 /i long x 330-415 fi wide) minutely punctured 
and with a few faint scattered reticulated areas (Text-fig. 55). " Anterior dorsal 
shield ' ' with twenty-one pairs of simple setae ; verticals well separated and directed 
anteriorly. " Posterior dorsal shield " with fifteen pairs of simple setae ; J5 very 




Figs. 55-58. Sejus laelaptoides (Berlese), female. Fig. 55, dorsum. Fig. 56, venter. 
Fig. 57, chelicera. Fig. 58, tarsus II. 



short. The distribution and relative lengths of the dorsal setae, and the ornamenta- 
tion of the dorsal shield are shown in the Text-fig. 

Tritosternum well developed with pilose laciniae. Sternal shield with a con- 
spicuous elliptical reticulated area, and bearing three pairs of simple setae (Text- 
fig. 56). Genital shield broad, posterior margin straight, and bearing one pair of 
simple setae. Ventri-anal shield large, broader than long (152-215 fi long x 240- 
291 /i wide), reticulated, and bearing eleven simple setae. Between the genital and 
ventri-anal shields lies a row of four narrow sclerotized plates. Stigma situated 
between coxae III and IV ; peritreme and peritrematal shield extending posterior 
to coxa IV. Metapodal plates small. 



THE PLATYSEIINAE (MESOSTIGM AT A : ACEOSEJIDAE) 55 

Venter of gnathosoma with rostral and internal palptrochanter setae long, whip- 
like. All setae on pedipalp simple. Tectum three-pronged and typical of the 
genus. The dentition of the chelicera is shown in Text-fig. 57. 

Leg I (536-673 fi long) with the tarsus (1 13-126 ju) longer than the tibia (104- 
118 /i). Setae on tarsus and tibia fine, those on other segments somewhat stouter ; 
tarsal claws strong. Tarsi II-IV with a pair of lanceolate setae ; ambulacra with 
three acuminate lobes (Text-fig. 58) . 

Male. Unknown. 

Distribution. Berlese (1887) described Epicrius laelaptoides from the Royal 
Botanic Gardens, Padua. Tragardh (1910) described Parasejus glaber var. minor 
from Sphagnum at Sarekgebirge, Sweden, and Halbert (1923) described Lasioseius 
(Episeius) sphagni from Sphagnum in Co. Dublin, Eire. The writers have examined 
Berlese's type in Florence, Halbert's type, and also numbers of females from Whern- 
side Moor, Cock Hill, and Malham Tarn, Yorkshire ; Rydal Water and Grasmere, 
Westmorland ; Leith Hill and Beckley, Surrey ; Lapland ; and Michigan. 



Sejus necorniger (Oudemans) comb. nov. 

Hypoaspis necorniger Oudemans, A. C, 1903, Ent. Ber., 1, 12 : 87. 

Platyseius necorniger, Buitendijk, A. M., 1945, Zool. Med., 24 : 296. 

Episeius necorniger, Willmann, C, 1949, Veroff. Mus. Nat. Bremen, No. iA : 120. 

Episeius necorniger, Willmann, C, 1952, Veroff. Inst. Meeresf. Bremerhaven, 1 : 148. 

Female. Dorsal shield (550-722 /i long x 320-467 /jl wide) completely covered 
by a fine reticulated pattern (Text-fig. 59). " Anterior dorsal shield " with twenty- 
one pairs of simple setae ; verticals, with their bases close together, considerably 
shorter than post- verticals. " Posterior dorsal shield " with fifteen pairs of simple 
setae. The distribution of the setae and the ornamentation of the dorsal shield are 
shown in the Text-fig. 

Tritosternum well developed with pilose laciniae. Sternal shield with some 
sculpturing, and with two semicircular marks anteriorly, and bearing three pairs of 
simple setae (Text-fig. 60). Genital shield convex posteriorly and bearing a single 
pair of setae. Ventri-anal shield slightly wider than long (162-190 /i long X 177- 
228 11 wide), reticulated, and bearing nine simple setae. Between the genital and 
ventri-anal shields lie four small sclerotized plates. Stigma situated between 
coxae III and IV ; peritreme extending posterior to coxa IV. Metapodal plates 
narrow. 

Venter of gnathosoma with rostral and internal palptrochanter setae long, whip- 
like. All setae on pedipalp simple. Tectum three-pronged and typical of the 
genus. The dentition of the chelicera is shown in Text-fig. 61. 

Leg I (578-782 pi long) with the tarsus (144-184 /i) longer than the tibia (101- 
126 /i) ; claws small, pul villus short. Setae on tarsus and tibia fine, those on 
remaining segments stouter. Tarsi II-IV with a pair of lanceolate setae ; ambulacra 
with three acuminate lobes (Text-fig. 62). 

Male. Unknown. 



56 



THE PLATYSEIINAE (MESOSTIGMATA : ACEOSEJIDAE; 



Distribution. Oudemans (1903) did not state the locality for his type female, 
but Buitendijk (1945) in his catalogue of the Oudemans Collection gives the locality 
as Borkum Island, Germany. Willmann (1952) records the species from the North 
Sea island of Wangerooge, and says it is found as far up as Lapland. We have 
examined specimens from the following localities : Kirkstone Pass, River Brathey 
and High Pike, Ambleside, Westmorland ; Rambulls Moor, Ilkley, Yorkshire ; 




Figs. 59-62. Sejus necorniger (Oudemans), female. Fig. 59, dorsum. Fig. 60, venter. 
Fig. 61, chelicera. Fig. 62, tarsus II. 



Hathersage, Derbyshire ; Sea Houses, Northumberland ; Burton Marsh, Cheshire ; 
Llyn Idwal, Caernarvonshire ; South Wales ; Washford, Somerset ; Kiel, Germany ; 
Ohrid, Montenegro ; and Grabouw, Transvaal. 



Sejus borealis (Berlese) comb. nov. 

Ameroseius borealis Berlese, A., 1904, Redia, 1 : 259. 

Episeius montanus Willmann, C, 1949, Veroff. Mus. Nat. Bremen, No. iA : 120. syn. nov. 
Episeius montanus Willmann, C, 1952, Veroff. Inst. Meeresf. Bremerhaven, 1 : 148. 
Episeius montanus Willmann, C, 1952, SB. ost. Akad. Wiss., Abt. 1, 162 : 462. 

Female. Dorsal shield (570-581 /i long x 361-371 /i wide) covered by a reticu- 
lated pattern (Text-fig. 63). " Anterior dorsal shield " bears twenty-one pairs of 
simple setae; verticals anteriorly projected. "Posterior dorsal shield" bears 



THE PLATYSEIINAE (MESOSTIGM AT A : ACEOSEJIDAE) 



57 



fifteen pairs of simple setae ; J5 are at the most half the length of J4. The majority 
of the dorsal setae arise from small tubercles. The distribution and relative length 
of the setae and the ornamentation of the dorsal shield are shown in the Text-fig. 
Tritosternum well developed and flanked at its base by a pair of narrow pre- 
endopodal shields. , Sternal shield with a narrow longitudinal reticulated area, and 
bearing three pairs of simple setae (Text-fig. 64). Genital shield convex posteriorly 
and bearing a single pair of setae. Ventri-anal shield broader than long (139-164 /i 
long X 202-205 ju, wide), reticulated and bearing eleven simple setae. Between the 
genital and ventri-anal shield lies a row of four small plates, and, posterior to these, 




Figs. 63-66. Sejus borealis (Berlese), female. Fig. 63, dorsum. Fig. 64, venter. 
Fig. 65, chelicera. Fig. 66, tarsus II. 



two smaller plates. Stigma situated between coxae III and IV with the posterior 
prolongation of the peritreme extending to about the middle of coxa IV. Metapodal 
plates circular. 

Venter of gnathosoma with rostral and internal palptrochanter setae long, whip- 
like. All setae on pedipalp simple. Tectum three-pronged and typical of the 
genus. The dentition of the chelicera is shown in Text-fig. 65. 

Leg I (576-580 /i long) with the tarsus (137-139 ji) longer than the tibia (108- 
114 ji). Setae on tarsus and tibia I fine ; ambulacrum and claws small ; remaining 
setae stouter and arising from small protuberances. Tarsi II-IV with a pair of 

ZOOL. 6, NO. 2. 5 



58 THE PLATYSEIINAE (MESOSTI GM ATA : ACEOSEJIDAE) 

lanceolate setae, and ambulacra with three acuminate lobes (Text-fig. 66). Setae 
of legs II-IV arising mainly from small protuberances. 

Male. Unknown. 

Distribution. Berlese's (1904) description of the female of Ameroseius borealis 
is based on material collected by Thor in Norway. His type specimen has been 
examined by us. Willmann (1949) described Episeius montanus from Poland ; in 
1952 he recorded the species from the North Sea island of Wangerooge, and in 1953 
he recorded it from several localities in the eastern Alps. The writers have 
examined five females from Soil Insecticide Experiment, Bellahouston Park, Glasgow, 
October, 1951, collected by J. G. Sheals ; one female from ditch-side debris near 
Markfield, Leicestershire, 19. iv. 1958, collected by P. N. Lawrence ; and one female 
from wet moss at West Carter Basin, Labrador, 17.viii.1958, also collected by 
P. N. Lawrence. 



Sejus serratus (Halbert) comb. nov. 

Paraseius serratus Halbert, J. N., 1915, Proc. R. Irish Acad., 39ii : 78. 
Lasioseius (L.) serratus, Schweizer, J., 1922, Ver. naturf. Ges. Basel, 33 : 43. 
Episeius serratus, Schweizer, J., 1949, Res. Rech. sci. Pare Nat. Suisse, N.F. 2 : 63. 
Episeius serratus, Willmann, C, 1949, Veroff. Mus. Bremen, 1A : 120 ; 1954, I n Spiegel, 
Land-Tierwelt, 15 Ordnung, Acarina : 351. 

Female. Dorsal shield (440-570 /i long x 286-351 /jl wide) with a pattern of 
irregular reticulations and depressions (Text-fig. 67). "Anterior dorsal shield" 
with twenty-one pairs of simple setae ; verticals on small tubercles and directed 
laterally. "Posterior dorsal shield" with fifteen pairs of simple setae. The 
distribution of the dorsal setae and the ornamentation of the dorsal shield are shown 
in the Text-fig. 

Tritosternum with a narrow base and pilose laciniae. Sternal shield with some 
sculpturing laterally, with a pair of circular marks anteriorly, and bearing three 
pairs of simple setae (Text-fig. 68). Genital shield convex posteriorly and bearing 
a single pair of setae. Ventri-anal shield slightly wider than long (152-170 p long 
X 165-200 }jl wide), reticulated anteriorly and bearing nine simple setae. Between 
the genital and ventri-anal shields lie a row of small plates and behind these a further 
pair. Stigma situated between coxae III and IV ; peritreme extending posterior to 
coxa IV. Metapodal plates small. 

Venter of gnathosoma with rostral and internal palptrochanter setae long, whip- 
like. All setae on pedipalp simple. Tectum three-pronged and typical of the genus. 
The dentition of the chelicera is shown in Text-fig. 69. 

Leg I (528-718 fi long) with the tarsus (121-149 /i) shorter than the tibia (139- 
164 fi). Claws on leg I small ; pulvillus short. Setae on tarsus and tibia fine ; 
those on remaining segments stouter. Tarsi II-IV with a pair of lanceolate setae ; 
ambulacra with three acuminate lobes. 

Male. This sex is described and figured by Schweizer (1922 & 1949). 

Distribution. Halbert (1915), in the Clare Island Survey, described this species 
from specimens " Found commonly in sphagnum, gathered on the slopes of Croagh- 



THE PLATYSEIINAE (MESOSTIGM AT A : ACEOSEJIDAE) 



59 



patrick at an elevation of about 600 ft., during the month of October ". Schweizer 
(1922 & 1949) records both sexes from the Swiss Alps, and Willmann (1954) records 
the female from Austria. The Museum Collections contain the following specimens : 
one female in Sphagnum from marsh, Raise Beck, Grasmere, Westmorland, 900 ft., 
29.vi.1951, collected by J. T. Salmon ; three females in mosses and liverworts on 
rocks by south-east bank of Rydal Water, Westmorland, 29 . vi . 1956, collected by 





Figs. 67-69. Sejus serratus (Halbert), female. Fig. 67, dorsum. Fig. 68, venter. 

Fig. 69, chelicera. 



P. N. Lawrence ; one female in Sphagnum at Blelham Tarn, Lancashire, 22 .iii. 1955, 
collected by R. J. Elliott ; one female in Sphagnum at Box Hill, Surrey, 30.vi.1951, 
collected by E. Duffey ; and two females in moss at West Carter Basin, Labrador, 
7.viii.i958, collected by P. N. Lawrence. 



Sejus curtipes (Halbert) comb. nov. 

Lasioseius (Episeius) glaber var. curtipes Halbert, J. N., 1923., /. Linn. Soc. Lond. (Zool.), 35 : 

370. 
Episeius ovaspini Schweizer, J., 1949, Res. Rech. sci. Pare Nat. Suisse, N.F. 2 : 69, syn. nov. 

Female. Dorsal shield (456-529 /i long x 258-320 /i wide) with a reticulated 
pattern (Text-fig. 70). " Anterior dorsal shield " with fifteen pairs of simple setae. 



6o 



THE PLATYSEIINAE (MESOSTIGM ATA : ACEOSEJIDAE; 



The distribution of the setae, and the ornamentation of the dorsal shield are shown 
in the Text-fig. 

Tritosternum with pilose laciniae. Sternal shield very faintly sculptured and 
bearing three pairs of simple setae (Text-fig. 71). Genital shield convex posteriorly 
and bearing a single pair of setae. Ventri-anal shield wider than long (120-160 /i 
long X 154-218 /i wide), reticulated in its anterior half, and bearing nine simple 
setae. Between the genital and ventri-anal shields lies a pair of long narrow 
platelets and behind these two shorter platelets. Stigma situated between coxae 
III and IV ; peritreme extending posterior to coxa IV. Metapodal plates small. 




Figs. 70-73. Sejus curtipes (Halbert), female. Fig. 70, dorsum. Fig. 71, venter. 
Fig. 72, chelicera. Fig. 73, tarsus II. 



Venter of gnathosoma with rostral and internal palptrochanter setae long, whip- 
like. All setae on pedipalp simple. Tectum three-pronged and typical of the 
genus. The dentition of the chelicera is shown in Text-fig. 72. 

Leg I (443-518 jlc long) with the tarsus (1 13-139 /i) longer than the tibia (68- 
88 jll). Claws on leg I small, pulvillus short ; setae on tarsus and tibia fine, those 
on remaining segments stouter. Tarsi II-IV with a pair of lanceolate setae ; 
ambulacra with three acuminate lobes (Text-fig. 73). 

Male. This sex is described and figured by Schweizer (1949 : 71). 

Distribution. Halbert's (1923) description of Lasioseius (Episeius) glaber var. 



THE PLATYSEIINAE (MESO STI GM AT A : ACEOSEJIDAE; 



6 i 



curtipes is based on two females found amongst wet moss on Lambay Island, Co. 
Dublin, in July. Schweizer's (1949) description of Episeius ovaspini is based on 
five males and thirty- two females collected in wet mosses in the Swiss National 
Park. We have examined Halbert's type and a number of Schweizer's type series ; 
also five females from the summit of Snowdon, Caernarvonshire, August, 195 1, 
collected by E. Duffey ; and eleven females from Jamaica, B.W.I., collected by 
P. F. Bellinger ; seven in moss on concrete, Coolsbade, Portland, 17.vii.1955, and 
four in moss, west side of John Cove Mountains, 2,300 ft., 25. v. 1956. 



Sejus cassiteridum sp. no v. 

Female. Dorsal shield (435-467 jlc long x 237-258 jlc wide) heavily sculptured 
and with a network of ridges and depressions. Lateral margins irregular (Text-fig. 
74). "Anterior dorsal shield" with twenty-one pairs of simple setae; verticals 




Figs. 74-78. Sejus cassiteridum sp. nov., female. Fig. 74, dorsum. Fig. 75, venter. 
Fig. 76, tectum. Fig. 77, chelicera. Fig. 78, tarsus II. 



close together and projecting anteriorly. "Posterior dorsal shield" with fifteen 
pairs of simple setae of which J5 are the shortest. The distribution and relative 
lengths of the dorsal setae and the ornamentation of the dorsal shield are shown in 
the Text-fig. 



62 THE PLATYSEIINAE (MESOSTIGM AT A : ACEOSEJIDAE) 

Tritosternum well developed with pilose laciniae. Sternal shield almost entirely 
without sculpturing, and bearing three pairs of short simple setae (Text-fig. 75). 
Genital shield slightly convex posteriorly and bearing a single pair of setae. Ventri- 
anal shield wider than long (1 15-139 /t long X 152-165 ji wide), reticulated and 
bearing nine simple setae. Between the genital and ventri-anal shields lie six 
small sclerotized plates. Stigma situated between coxae III and IV ; peritreme 
and peritrematal shields extending posterior to coxa IV. Metapodal plates small. 

Venter of gnathosoma with rostral and internal palptrochanter setae long, whip- 
like. All setae on pedipalp simple. Tectum (Text-fig. 76) three-pronged. The 
dentition of the chelicera is shown in Text-fig. yy. 

Leg I (372-387 ju, long) with the tarsus (99-106 /i) almost twice the length of the 
tibia (58-63 fi). Setae on tarsus and tibia fine, those on other segments stouter ; 
pulvillus short and tarsal claws small. Tarsi II-IV with a pair of lanceolate setae ; 
ambulacra with three acuminate lobes (Text-fig. 78). 

Male. Unknown. 

Locality. England. The holotype female (1959. 1.20. 92) and twenty-five 
paratype females (1959 . 1 . 20 . 93-102) occurring with Platyseius subglaber (Oudemans) 
in the roots of rushes by Big Pool, St. Agnes, Isles of Scilly, Cornwall, 19. ix. 1957, 
collected by K. H. Hyatt. 

The writers have examined a single female (1959. 1.20. 193) from Sphagnum, 
Blelham Tarn, Windermere, Lancashire, 22 .iii. 1955, collected by R. J. Elliott, which 
differs from the St. Agnes specimens by having the sternal shield conspicuously 
reticulated. The measurements of this specimen all fall within the ranges given 
above for Sejus cassiteridum (viz. : dorsal shield 446 X 247 jlc, ventri-anal shield 
126 X 157 fi, leg I 392 /i, tarsus I 101 /i, tibia I 63 /i). 



Sejus dromadis sp. nov. 

Female. Dorsal shield (393 /i long x 230 pu wide) oval in outline, heavily 
reticulated, and steeply humped dorso-medially (Text-fig. 79). " Anterior dorsal 
shield " with twenty- three pairs of setae ; " posterior dorsal shield " with fifteen 
pairs of setae (Text-fig. 80). The verticals are stout and spiculate, and the re- 
mainder of the setae are stout and pilose. The distribution and relative lengths of 
the dorsal setae, and the form of the dorsal shield are shown in the Text-figs. 

Tritosternum less strongly developed than in the majority of species of Sejus. 
Laciniae pilose. Sternal shield weakly sclerotized with three pairs of simple setae. 
Genital shield broadly flask-shaped with one pair of setae (Text-fig. 81). Ventri- 
anal shield small (83 ju long X 88 jll wide), weakly sclerotized, and bearing seven 
simple setae. Two small sclerotized plates lie on the interscutal membrane between 
the genital and ventri-anal shields. Stigma situated between coxae III and IV, 
with the peritreme not extending posterior to the stigma. Peritrematal shield 
extending a little posterior to the stigma. 

Venter of gnathosoma with rostral and internal palptrochanter setae long and 
whip-like (Text-fig. 82). All setae on pedipalp simple. Tectum three-pronged and 



THE PLATYSEIINAE (MESOSTIGM AT A : ACEOSEJIDAE) 63 




Figs. 79-85. Sejus dromadis sp. nov., female. Fig. 79, lateral view of dorsum. Fig. 
80, dorsal shield. Fig. 81, venter. Fig. 82, venter of gnathosoma. Fig. 83, tectum. 
Fig. 84, chelicera. Fig. 85, tarsus II. 



6 4 



THE PLATYSEIINAE (MESOSTI GM AT A : ACEOSEJIDAE) 



typical of the genus (Text-fig. 83). The dentition of the chelicera is shown in 
Text-fig. 84. 

Leg I (about 300 pi long) with the tarsus (70 fi) longer than the tibia (56 jlc). Setae 
on tarsus fine ; those on remaining segments stouter, and in some cases spiculate 
and arising from small protuberances. Tarsi II-IV with a pair of lanceolate setae. 
Ambulacrum of tarsus I short and claws small. Pulvilli of tarsi II-IV produced 
into three acuminate lobes (Text-fig. 85). 

Male. Unknown. 

Locality. England. A single female (1959. 1.20. 103) in cow dung at Canter- 
bury, Kent, collected by the late Dr. E. Warren. 



Sejus areolatus sp. nov. 

Female. Dorsal shield (361 /i long x 216 /i wide) completely covered by a fine 
reticulated pattern; margin irregular (Text-fig. 86). "Anterior dorsal shield" 




Figs. 86-90. Sejus areolatus sp. nov., female. Fig. 86, dorsum. Fig. 87, venter. Fig. 88, 
tectum. Fig. 89, chelicera. Fig. 90, tarsus II. 



with twenty-one pairs of simple setae ; verticals with their bases touching, and 
considerably shorter than post- verticals. "Posterior dorsal shield" with fifteen 
pairs of simple setae. The distribution of the setae and the ornamentation of the 
dorsal shield are shown in the Text-fig. 

Tritosternum well developed with pilose laciniae. Sternal shield almost plain, 



THE PLATYSEIINAE (MESOSTIGM AT A : ACEOSEJIDAE) 65 

with three pairs of simple setae (Text-fig. 8y). Genital shield bearing a single pair 
of setae. Ventri-anal shield wider than long (106 /i long x 132 fi wide), reticulated, 
with five small oval areas forming a crescent behind the paranals, and bearing nine 
simple setae. Between the genital and ventri-anal shields lies a row of four sclero- 
tized platelets, and behind these two smaller ones. Stigma situated between coxae 
III and IV ; peritreme extending posterior to coxa IV. Metapodal plates small and 
narrow. 

Venter of gnathosoma with rostral and internal palptrochanter setae long, whip- 
like. All setae on pedipalp simple. Tectum three-pronged (Text-fig. 88), the centre 
prong being in advance of the lateral ones. The dentition of the chelicera is shown 
in Text-fig. 89. 

Leg I (c. 330 /i long) with the tarsus (88 /i) longer than the tibia (50 /£). Setae 
on tarsus and tibia fine, those on remaining segments stouter ; claws small, on short 
pul villus. Tarsi II-IV with a pair of lanceolate setae ; ambulacra with three 
acuminate lobes (Text-fig. 90). 

Male. Unknown. 

Locality. Uganda. A single female, the holotype (1959. 1.20. 104), from 
rushes and grassland bordering hot springs, sixteen miles east-north-east of Bundi- 
bugyo, Ruwenzori, 29.viii.1952, collected by G. Owen Evans. 



Sejus kennedyi sp. nov. 

Female. Dorsal shield (456-467 /i long x 268-279 /i wide) covered by a 
network of small depressions and ridges (Text-fig. 91). " Anterior dorsal shield" 
with twenty-one pairs of simple setae ; verticals separated by the width of their 
bases and projecting anteriorly. " Posterior dorsal shield" with fifteen pairs of 
simple setae of which J5 are short. The distribution and relative lengths of the 
dorsal setae and the ornamentation of the dorsal shield are shown in the Text-fig. 

Tritosternum with a pair of pilose laciniae. Sternal shield plain and bearing 
three pairs of simple setae (Text-fig. 92). Genital shield lightly reticulated, convex 
posteriorly, and bearing one pair of simple setae. Ventri-anal shield broader than 
long (139-144 /i long x 208-213 jlc wide), reticulated, and bearing nine simple setae. 
Between the genital and ventri-anal shields lie two narrow sclerotized platelets. 
Stigma situated between coxae III and IV ; peritreme crenate along its internal 
margin, and with no post-stigmal prolongation. Peritrematal shield tapering and 
extending posterior to coxa IV. Metapodal shields obliquely situated. 

Venter of gnathosoma with rostral and internal palptrochanter setae long, whip- 
like. All setae on pedipalp simple. Tectum three-pronged and typical of the 
genus. Chelicerae typical. 

Leg I (354 ju, long) with the tarsus (88 fi) longer than the tibia (63 /i) . Setae on 
tarsus I fine, ambulacrum and claws small ; setae on remaining segments stouter 
and arising from small protuberances. Tarsi II-IV with a pair of lanceolate setae 
and ambulacra with three acuminate lobes. 

Male. Unknown. 



66 



THE PLATYSEIINAE (MESOSTIGM AT A : ACEOSEJIDAE] 



Locality. Uganda. The holotype female (1959 . 1 . 20 . 105) and three paratype 
females (1959 . 1 . 20 . 106-108) from litter under hardwood stand, Bundibugyo, 
Ruwenzori, 3,400 ft., 2.ix.i952, collected by G. Owen Evans. This species is 
named after Professor W. Q. Kennedy, University of Leeds, leader of the British 
Ruwenzori Expedition, 1952. 





92 
91 

Figs. 91-92. Sejus kennedyi sp. nov., female. Fig. 91, dorsum. Fig. 92, venter. 



Sejus phalangioides sp. nov. 

Female. Dorsal shield (398-410 fi long x 250-253 fi wide) attenuated in the 
posterior half, and covered by a fine reticulated pattern (Text-fig. 93). " Anterior 
dorsal shield " with twenty-one pairs of simple setae, verticals projecting anteriorly ; 
" posterior dorsal shield" with fifteen pairs of simple setae, of which J5 are the 
shortest. The distribution and relative lengths of the dorsal setae are shown in 
the Text-fig. 

Tritosternum well developed and flanked at its base by a pair of narrow pre- 
endopodal shields. Sternal shield with a reticulated area anteriorly, and bearing 
three pairs of simple setae (Text-fig. 94) . Genital shield broad, truncated posteriorly, 



THE PLATYSEIINAE (MESOSTIGM AT A : ACEOSEJIDAE) 



67 



distinctly reticulated, and bearing a single pair of setae. Ventri-anal shield slightly 
broader than long (101 /i long X 116 ja wide), attenuated posteriorly, reticulated, 
and bearing nine simple setae. Between the genital and ventri-anal shields lies a 
narrow, undulated, sclerotized strip of chitin, and two platelets. Stigma situated 
between coxae III and IV, peritreme scarcely extending posterior to the stigma. 




Figs. 93-99. Sejus phalangioides sp. nov., female. Fig. 93, dorsum. Fig. 94, venter. 
Fig. 95, tectum. Fig. 96, chelicera. Fig. 97, leg IV. Fig. 98, tarsus II. Fig. 99, 
entire mite, showing relative lengths of legs and idiosoma. 



Peritrematal shield fused with well developed podal plates, which extend posterior 
to coxa IV. Metapodal plates small, situated close to podal plates. 

Venter of gnathosoma with rostral and internal palptrochanter setae long, whip- 



68 



THE PLATYSEIINAE (MESOSTIGMATA : ACEOSEJIDAE) 



like. All setae on pedipalp simple. Tectum (Text-fig. 95) three-pronged and 
typical of the genus. The dentition of the chelicera is shown in Text-fig. 96. 

Leg I (536 jlc long) has the tarsus (139 ja) longer than the tibia (100-106 jll). Setae 
on tarsus I very fine ; those on remaining segments stouter. Ambulacrum of 
tarsus I short, claws small. Leg IV (954 /i long) more than twice the length of the 
idiosoma, with the tarsus about 400 pi long (Text-fig. 97). Tarsi II— III with a pair 
of lanceolate setae, and pulvilli produced into three acuminate lobes (Text-fig. 98). 
Tarsus IV without lanceolate setae, and pretarsus long (55 /i). The relative lengths 
of the legs and the idiosoma are shown in Text-fig. 99. 

male. Unknown. 

Locality. Uganda. The holotype female (1959. 1.20. 109) and one paratype 
female (1959. 1.20. no) in grassland in forest clearing near stream, nine miles 
north-east of Bundibugyo, Ruwenzori, 2,850 ft., 24.viii.1952, collected by G. Owen 
Evans. 



Sejus signatus sp. nov. 

Female. Dorsal shield (581 [i long X 371 fi wide) weakly sclerotized and covered 
by a reticulated pattern (Text-fig. 100). "Anterior dorsal shield" with twenty 




Figs. 100-102. Sejus signatus sp. nov., female. Fig. 100, dorsum. Fig. 101, venter. 

Fig. 102, chelicera. 



THE PLATYSEIINAE (ME SOSTI GM AT A : ACEOSEJIDAE) 69 

pairs of simple setae ; verticals considerably shorter than post-verticals, and well 
separated. "Posterior dorsal shield" with fifteen pairs of simple setae. The 
distribution of the dorsal setae and the ornamentation of the dorsal shield are shown 
in the Text-fig. 

Tritosternum with pilose laciniae. Sternal shield weakly sclerotized, with an 
M-shaped mark anteriorly and bearing three pairs of simple setae (Text-fig. 101). 
Genital shield weakly sclerotized and bearing one pair of setae. Ventri-anal shield 
wider than long (152 /i long X 197 /i wide), weakly sclerotized and bearing nine 
simple setae. Between the genital and ventri-anal shields lie a narrow sclerotized 
strip and two platelets. Stigma situated between coxae III and IV ; peritreme 
extending posterior to coxa IV. Metapodal plates very small. 

Venter of gnathosoma with rostral and internal palptrochanter setae long, whip- 
like. All setae on pedipalp simple. Tectum three-pronged and typical of the 
genus. The dentition of the chelicera is shown in Text-fig. 102. 

Leg I (573 jlc long) with the tarsus (141 /i) longer than the tibia (99 fi). Claws 
on leg I small, inconspicuous, pulvillus short. Setae on tarsus and tibia fine, those 
on other segments stouter. Tarsi II-IV with a pair of lanceolate setae ; ambulacra 
with three acuminate lobes. 

Male. Unknown. 

Locality. Uganda. A single female, the holotype (1959.1.20.111), from damp 
decaying vegetation and soil under elephant-grass at Ibanda, Ruwenzori, 26 . vii . 1952, 
4,760 ft., collected by G. O. Evans. 



Sejus spinipes sp. nov. 

Female. Dorsal shield (392 pi long X 216 pi wide) covered by a light reticulated 
pattern (Text-fig. 103). " Anterior dorsal shield " bears twenty-one pairs of simple 
setae ; verticals anteriorly projecting. " Posterior dorsal shield" bears 15 pairs of 
simple setae ; J5 are as long as J4. The majority of the dorsal setae arise from small 
tubercles. The distribution and relative lengths of the dorsal setae, and the orna- 
mentation of the dorsal shield are shown in the Text-fig. 

Tritosternum well developed with pilose laciniae. Sternal shield with the central 
area reticulated, and bearing three pairs of short simple setae (Text-fig. 104). 
Genital shield with a single pair of simple setae. Ventri-anal shield broader than 
long (119 /i long x 157 jlc wide), distinctly reticulated and bearing nine simple 
setae. Between the genital and ventri-anal shields lies a line of four contiguous 
sclerotized plates, and, postero-laterally to these, four smaller plates. Stigma 
situated between coxae III and IV, with the posterior prolongation of the peritreme 
extending to about the middle of coxa IV. The peritrematal shield extends posterior 
to coxa IV. Metapodal plates small. 

Venter of gnathosoma with rostral and internal palptrochanter setae long, whip- 
like. All setae on pedipalp simple. Tectum (Text-fig. 105) three-pronged. The 
dentition of the chelicera is typical of the genus. 



7 o 



THE PLATYSEIINAE (MESOSTIGM AT A : ACEOSEJIDAE 



Leg I ( c. 330 pi long) with the tarsus (76 ft) longer than the tibia (53 fi). Setae 
on tarsus I fine ; ambulacrum and claws small ; many of the setae on remaining 
segments stout and arising from conspicuous protuberances. Tarsi II-IV with a 
pair of lanceolate setae, and ambulacra with three acuminate lobes. Setae of legs 
II-IV mainly stout and arising from conspicuous protuberances. 

Male. Unknown. 

Localities. Uganda and Sudan. The holotype female (1959. 1.20. 112) from 




Figs. 103-105. Sejus spinipes sp. nov., female. Fig. 103, dorsal shield. Fig. 104, venter. 

Fig. 105, chelicera. 



rushes and grassland bordering hot springs, sixteen miles east-north-east of Bundi- 
bugyo, Ruwenzori, Uganda, 29.viii.1952, collected by G. O. Evans; and two 
paratype females (1959.1.20.113-114) on Papyrus in the Sudd Region, Southern 
Sudan, collected by I. W. B. Thornton. 



THE PLATYSEIINAE (MESOSTIGM AT A : ACEOSEJIDAE) 71 

Sejus nepalensis sp. nov. 

Female. Dorsal shield (456-498 ju, long x 289-299 ju, wide) completely covered 
by a fine reticulated pattern (Text-fig. 106). " Anterior dorsal shield " with twenty- 
one pairs of simple setae ; vertical setae relatively short and directed laterally ; 
vertex not well developed. " Posterior dorsal shield " with fifteen pairs of simple 
setae (in the holotype seta Zi is missing from the left side) ; seta J5 is the shortest. 
All dorsal setae arise from small tubercles. The distribution and relative lengths of 
the setae and the ornamentation of the dorsal shield are shown in the Text-fig. 

Tritosternum well developed, with pilose laciniae, sternal shield with some 




Figs. 106-109. Sejus nepalensis sp. nov., female. Fig. 106, dorsum. Fig. 107, venter. 
Fig. 108, chelicera. Fig. 109, tarsus II. 



sculpturing, a pair of circular areas anteriorly, and bearing three pairs of simple 
setae (Text-fig. 107). Genital shield almost parallel sided and bearing one pair of 
simple setae. Ventri-anal shield (126 /jl long X 124-127 jlc wide) almost circular in 
outline, bearing nine simple setae ; anus more or less central. Between the genital 
and ventri-anal shields lie about four narrow platelets. Stigma situated between 
coxae III and IV ; peritreme extending posterior to coxa IV. Metapodal plates 
very small. 

Venter of gnathosoma with rostral and internal palptrochanter setae long, whip- 
like. All setae on pedipalp simple. Tectum three-pronged and typical of the 
genus. The dentition of the chelicera is shown in Text-fig. 108. 



7 2 



THE PLATYSEIINAE (MESOSTIGM AT A : ACEOSEJIDAE) 



Leg I (630-769 /i long) with the tarsus (126-157 /i) shorter than the tibia (150- 
195 /£). Setae on tarsus, tibia and genu very fine ; those on remaining segments 
stouter. Claws very small, pul villus short. Tarsi II-IV with a pair of lanceolate 
setae ; ambulacra with three acuminate lobes (Text-fig. 109). 

Male. Unknown. 

Locality. Nepal. The holotype female (1959. 1.20. 115) from sandy turf on 
open grazed hillside facing south at Siklis (28 22' N., 84 6' E.), 7,000 ft., 20- 
21. iv. 1954; and one paratype female (1959. 1.20. 116) from damp mossy and 
grassy earth under shade of trees and rocks at Bakhri Kharka (28 22-5' N., 84 
7-5' E.), 5,500 ft., 24. iv. 1954, collected by K. H. Hyatt. 



Sejus parbatensis sp. nov. 

Female. Dorsal shield (402-435 fi long x 237-268 jlc wide) covered by an 
irregular reticulated pattern (Text-fig. no). " Anterior dorsal shield " with twenty- 




Figs. 110-113 



Sejus parbatensis sp. nov., female. Fig. no, dorsum. 
Fig. 112, chelicera. Fig. 113, tarsus II. 



one pairs of simple setae ; verticals with their bases contiguous. " Posterior 
dorsal shield ' ' with fifteen pairs of simple setae ; setae J5 are more than half the 
length of J4. All the setae arise from small tubercles. The distribution of the 
setae, and the ornamentation of the dorsal shield are shown in the Text-fig. 



THE PLATYSEIINAE (MESOSTIGM AT A : ACEOSEJIDAE) 73 

Tritosternum well developed and with pilose laciniae. Sternal shield lightly 
sculptured and bearing three pairs of simple setae (Text-fig. in). Genital shield 
bearing a single pair of setae. Ventri-anal shield wider than long (1 14-126 jlc long 
X 157-170 fi wide), reticulated, and bearing nine simple setae. Between the genital 
and ventri-anal shields lies a row of four narrow platelets, and, posterior to these, 
two smaller ones. Stigma situated between coxae III and IV ; peritreme extending 
posterior to coxa IV. Metapodal plates very small. 

Venter of gnathosoma with rostral and internal palptrochanter setae long, whip- 
like. All setae on pedipalp simple. Tectum three-pronged and typical of the 
genus. The dentition of the chelicera is shown in Text-fig. 112. 

Leg I (c. 340 /i long) with the tarsus (80-83 M>) longer than the tibia (50-53 /i). 
Setae on tarsus I very fine ; those on remaining segments stouter ; claws small, 
on short pulvillus. Tarsi II-IV with a pair of lanceolate setae ; ambulacra with 
three acuminate lobes (Text-fig. 113). 

Male. Unknown. 

Locality. Nepal. The holotype female (1959. 1.20. 117) and one paratype 
(1959. 1. 20. 1 18) from litter under a group of deciduous trees forming a canopy on 
open ground at Gurjakhani (28 36-5' N., 83 13-5' E.) on the southern slope of the 
Dhaulagiri Himal, 8,500 ft., 24-27 . vi . 1954 ; and one paratype female (1959. 1.20. 
119) in sandy turf on open, grazed hillside facing south at Siklis (28 22' N., 84 6' 
E.), 7,000 ft., 20-21. iv. 1954, collected by K. H. Hyatt. 



Sejus clayi sp. nov. 

Female. Dorsal shield (661 /i long x 366 /i wide) with lateral incisions anterior 
to Si, and heavily marked with circular depressions (Text-fig. 114). Vertex of 
dorsal shield produced anteriorly (Text-fig. 115). "Anterior dorsal shield" with 
twenty pairs of setae ; verticals absent, para- verticals short and spatulate, remaining 
setae simple. " Posterior dorsal shield " with fifteen pairs of simple setae. Setae 
J5 are long. The distribution and relative lengths of the dorsal setae are shown in 
Text-fig. 114. 

Tritosternum well developed with pilose laciniae and flanked at its base by a 
number of fragmented pre-endopodal shields. Sternal shield plain and bearing 
three pairs of simple setae (Text-fig. 116). Genital shield wedge shaped, genital 
setae off the shield. Ventri-anal shield slightly wider than long (185 jlc long X 190 /i 
wide), covered with small circular depressions, and bearing only five simple setae. 
Stigma situated between coxae III and IV, peritreme not extending posterior to the 
stigma ; external margin of peritreme strongly crenate. Peritrematal shield 
extending posterior to coxa IV. Metapodal shields narrow and obliquely situated. 

Venter of the gnathosoma with rostral and internal palptrochanter setae long, 
whip-like. All setae on legs simple. Tectum (Text-fig. 117) differing from that of 
other members of the genus in that the median prong is strongly produced and 
terminates in a pair of pilose laciniae. The dentition of the chelicera is shown in 
Text-fig. 118. 

ZOOL. 6, NO. 2. 6 



74 



THE PLATYSEIINAE (MESOSTIGM AT A : ACEOSEJIDAE 



Leg I (481 /i long) with the tarsus (126 fi) longer than the tibia (101 fi). Setae 
on tarsus I very fine ; those on remaining segments stouter. Tarsi II-IV with a 
pair of lanceolate setae and ambulacra with lateral lobes acuminate, median lobe 
short and rounded (Text-fig. 119). Ambulacrum of tarsus I short with claws very 
small. 

Male. Unknown. 









118 

Figs. 114-119. Sejus clayi sp. nov., female. Fig. 114, dorsum. Fig. 115, vertex. 
Fig. 116, venter. Fig. 117, tectum. Fig. 118, chelicera. Fig. 119, tarsus II. 

Locality. Sikkim. A single female (1959. 1.20. 120) from leaves and leaf- 
mould in wood on steep hillside, Chungtang, 5,120 ft., 16. ii. 1952, collected by 
Dr. Theresa Clay. 



Sejus alstoni sp. nov. 

Female. Dorsal shield (507-550 jlc long x 279-340 jlo wide) covered anteriorly 
by a network of small depressions and ridges, and posteriorly by a reticulated 



THE PLATYSEIINAE (MESOSTIGM AT A : ACEOSEJIDAE) 



75 



pattern (Text-fig. 120). " Anterior dorsal shield " with twenty-one pairs of simple 
setae ; verticals close together and directed anteriorly. " Posterior dorsal shield " 
with fifteen pairs of simple setae, of which J5 are the shortest. The distribution 
and relative lengths of the dorsal setae, and the ornamentation of the dorsal shield 
are shown in the Text-fig. 




Figs. 120-123. Sejus alstoni sp. nov., female. Fig. 120, dorsum. 
Fig. 122, tectum. Fig. 123, tarsus II. 



Tritosternum well developed with pilose laciniae, and flanked at its base by a 
pair of narrow pre-endopodal shields. Sternal shield weakly sclerotized, without 
ornamentation, and bearing three pairs of simple setae (Text-fig. 121). Genital 
shield truncated posteriorly and bearing one pair of setae. Ventri-anal shield 
broader than long (139-152 fx long X 180-205 fi wide), reticulated, and bearing nine 
simple setae. Between the genital and ventri-anal shields lies a row of three narrow 
sclerotized plates. Stigma situated between coxae III and IV ; no posterior 
prolongation of the peritreme. Peritrematal shield extending posterior to coxa IV. 
Metapodal shields elliptical. 

Venter of gnathosoma with rostral and internal palptrochanter setae long, whip- 



76 THE PLATYSEIINAE (MESOSTIGMATA : ACEOSEJIDAE) 

like. All setae on pedipalp simple. Tectum (Text-fig. 122) three-pronged and 
typical of the genus. Chelicerae typical. 

Leg I (560-565 ji long) with the tarsus (160 /i) one-and-a-half times as long as the 
tibia (101-104 /i). Setae on tarsus fine ; claws small ; setae on remaining segments 
stouter. Tarsi II-IV with a pair of lanceolate setae and ambulacra with three 
acuminate lobes (Text-fig. 123). 

Male. Unknown. 

Locality. Indonesia. The holotype female (1959. 1.20. 121) and two paratype 
females (1959. 1.20. 122-123) in decaying vegetation at Bogor, 1954, collected by 
the late A. H. G. Alston. 



Sejus aciculatus sp. nov. 

Female. Dorsal shield (329-341 /i long x 223-231 fi wide) very heavily sclero- 
tized, entirely covered by a network of ridges and depressions (Text-fig. 124). 
" Anterior dorsal shield" with nineteen pairs of simple setae, verticals and para- 
verticals apparently absent. " Posterior dorsal shield " with fifteen pairs of simple 
setae. The distribution and relative lengths of the dorsal setae, and the charac- 
teristic ornamentation of the dorsal shield are shown in the Text-fig. 

Tritosternum well developed and flanked at its base by a pair of strong pre- 
endopodal shields which abut the sternal shield. Sternal shield characteristically 
ornate (Text-fig. 125) and bearing three pairs of simple setae. Genital shield 
approximately in the form of a trapezium, heavily sclerotized and ornate, and bearing 
a single pair of simple setae. Ventri-anal shield broader than long (88-96 ju, long 
X 172-180 pt wide), concave anteriorly, strongly ornate, and bearing nine simple 
setae. Between the genital and ventri-anal shields lies a pair of narrow sclerotized 
platelets. Stigma situated between coxae III and IV and peritreme not extending 
posterior to the stigma. Peritrematal shield well developed and extending posterior 
to coxa IV. Metapodal plates small. 

Venter of gnathosoma with rostral and internal palptrochanter setae long, whip- 
like. All setae on pedipalp simple. Tectum (Text-fig. 126) three-pronged, median 
prong broad and triangular, lateral prongs small, inconspicuous. Dentition of 
chelicera as in Text-fig. 127. 

Leg I (245-255 /i long) with the tarsus (88-94 fi) twice the length of the tibia 
(43-45 /i). Setae on tarsus I very fine, ambulacrum short, claws small. Setae of 
remaining segments stouter. Tarsi II-IV with a pair of lanceolate setae, and 
ambulacra with three acuminate lobes (Text-fig. 128). 

Male. Dorsal shield (276-283 /i long x 180-185 /* wide) with ornamentation and 
chaetotaxy similar to the female. 

Tritosternum of the same form as that of the female. The chaetotaxy and 
sclerotization of the venter are shown in Text-fig. 129. The gnathosoma and 
tectum are similar to those of the female. The spermatophoral process is about the 
same length as the movable digit of the chelicera (Text-fig. 130). 



THE PLATYSEIINAE (MESOSTIGM AT A : ACEOSEJIDAE) 



77 




Figs. 124-130. Sejus aciculatus sp. nov. Fig., 124 dorsal shield of female. Fig. 125, 
venter of female. Fig. 126, tectum of female. Fig. 127, chelicera of female. Fig. 128, 
tarsus II of female. Fig. 129, venter of male. Fig. 130, chelicera of male. 



78 THE PLATYSEIINAE (MESOSTIGMATA : ACEOSEJIDAE) 

Leg I (281 ja long) with the tarsus (81 jlc) twice the length of the tibia (38 jlc). The 
chaetotaxy and ambulacra of the legs are as in the female. 

Locality. Jamaica, B.W.I. The holotype female (1959. 1.20. 124), allotype 
male (1959. 1.20. 125), seven female and three male paratypes (1959. 1.20. 126-133) 
from damp leaf litter at St. Ann, Mt. Diablo, 1,500 ft. and 2,200 ft., 3.VL1956, 
collected by P. F. Bellinger. A male and female paratype have been returned to 
the collector. 



Sejus antillanus sp. nov. 

Female. Dorsal shield (333-443 /i long x 195-278 jlc wide) slightly attenuated 
in its posterior half, and entirely covered by a reticulated pattern (Text-fig. 131). 
"Anterior dorsal shield" with twenty-one pairs of simple setae, verticals almost 
contiguous and directed anteriorly. " Posterior dorsal shield" with fifteen pairs 
of simple setae ; J5 are the shortest. The distribution and relative lengths of the 
setae, and the ornamentation of the dorsal shield are shown in the Text-fig. 

Tritosternum well developed with pilose laciniae. Sternal shield with a reticulated 
area anteriorly and bearing three pairs of simple setae (Text-fig. 132). Genital 
shield truncated posteriorly, reticulated, and bearing a single pair of setae. Ventri- 
anal shield broader than long (76-101 jli long x 101-114 jli wide), reticulated, with 
nine simple setae of which one pair is long. Between the genital and ventri-anal 
shields lies a row of four small narrow sclerotized plates, and behind these a pair 
of smaller plates. Stigma situated between coxae III and IV ; no post-stigmal 
prolongation of the peritreme. Peritrematal shield extending posterior to coxae IV. 
Metapodal shields small. 

Venter of gnathosoma with rostral and internal palptrochanter setae long, whip- 
like. All setae on pedipalp simple. Tectum (Text-fig. 133) three-pronged. The 
dentition of the chelicera is shown in Text-fig. 134. 

Leg I (384-521 jli long) with the tarsus (86-139 jli) slightly longer than the tibia 
(78-114 jlc). Setae on tarsus and tibia fine, those on remaining segments stouter ; 
claws on leg I small. Tarsi II-IV with a pair of lanceolate setae and ambulacra 
with acuminate lobes (Text-fig. 135). Leg IV (532^797 jlc) considerably longer than 
the idiosoma, with the tarsus 203-316 jli long. 

Male. Dorsal shield (236-303 jlc long x 147-203 jlc wide) having similar chaeto- 
taxy and ornamentation to the female. 

The chaetotaxy and sclerotization of the venter are shown in Text-fig. 136. The 
gnathosoma and tectum are similar to those of the female. The spermatophoral 
process is about one-and-a-half times the length of the movable digit (Text-fig. 137) . 

Leg I (238-432 jlc long) with the tarsus (76-116 jli) longer than the tibia (63-88 jlc). 
Leg IV (448-658 jlc long) with the tarsus 170-253 jlc. The chaetotaxy and ambulacra 
of the legs are as in the female. 

Locality. Jamaica, B.W.I. The holotype female (1959. 1.20. 134) and four 
paratype females (1959. 1.20. 135-138) from damp leaf litter, Mt. Diablo, St. Ann, 
2,200 ft., 7.VL1956 ; the allotype male (1959. 1.20. 139) and five paratypes (3<J(J, 



THE PLATYSEIINAE (MESOSTIGM AT A : ACEOSEJIDAE) 



79 




Figs. 131-137. Sejus antillanus sp. nov. Fig. 131, dorsum of female. Fig. 132, venter 
of female. Fig. 133, tectum of female. Fig. 134, chelicera of female. Fig. 135, 
tarsus II of female. Fig. 136, venter of male. Fig. 137, chelicera of male. 



8o 



THE PLATYSEIINAE (MESOSTIGM AT A : ACEOSEJIDAE) 



2 ??) (1959 . 1 . 20 . 140-144) from damp leaf litter in wood, Dolphin Head, Hanover, 
1,780 ft., 13. iv. 1956 ; three paratypes (ig, 2$?) in moss, St. Thomas, John Cove 
Mountains, 2,300 ft., 25. v. 1956 ; and ten paratypes (i<J, 9$$) (1959. 1.20. 145-154) 
from leaf litter, Portland Gap, 8. hi. 1956. All were collected by P. F. Bellinger. 
A male and two female paratypes have been returned to the collector. 




Figs. 138-142. Sejus bellingeri sp. nov. Fig. 138, dorsum of female. Fig. 139, tectum 
of female. Fig. 140, tarsus I of female. Fig. 141, venter of male. Fig. 142, chelicera 
of male. 



THE PLATYSEIINAE (MESOSTIGM AT A : ACEOSEJIDAE) 81 

Sejus bellingeri sp. nov. 

Female. Dorsal shield (467-507 /i long x 299-310 p wide) slightly attenuated 
in its posterior half, and covered by a reticulated pattern (Text-fig. 138). " Anterior 
dorsal shield ' ' with twenty-one pairs of simple setae ; verticals closely situated and 
directed anteriorly. " Posterior dorsal shield " with fifteen pairs of simple setae of 
which J5 are the shortest. The distribution and relative lengths of the setae and 
the ornamentation of the dorsal shield are shown in the Text-fig. 

The chaetotaxy and sclerotization of the venter are essentially the same as in Sejus 
nodosus (p. 85). The ventri-anal shield measures 119-139 p, long X 132-156 fi 
wide. 

Venter of gnathosoma with rostral and internal palptrochanter setae long, whip- 
like. All setae on pedipalp simple. Tectum (Text-fig. 139) three-pronged, and 
chelicerae typical of the genus. 

Leg I (c. 589 p long) with the tarsus (c. 157 p) longer than the tibia (c. 126 p). 
Tarsus I with a strong erect seta dorsally (Text-fig. 140) , remaining setae fine ; 
tarsal claws small but conspicuous. Tarsi II— III with a pair of lanceolate setae 
(tarsus IV has only one such seta), ambulacra with three acuminate lobes. Leg IV 
(c. 913 pi) longer than the idiosoma. Tarsus IV (303-341 fi) markedly elongate and 
three times the length of the tibia. 

Male. Dorsal shield (347 pi long X 228 pi wide) not attenuated in its posterior 
half, lightly reticulated. The chaetotaxy of the dorsal shield is the same as that of 
the female. 

The chaetotaxy and ornamentation of the venter are shown in Text-fig. 141. 
The gnathosoma and tectum are similar to those of the female. Spermatophoral 
process about twice the length of the movable digit of the chelicera (Text-fig. 142). 

Leg I (481 pi long) with the tarsus (126 p) longer than the tibia (99 p). Tarsus I 
with a strong erect seta dorsally as in the female. Leg IV (675 p long) longer than 
the idiosoma, and with the tarsus (265 p) three times the length of the tibia (88 p). 

Locality. Jamaica, B.W.I. The holotype female (1959. 1.20. 155), allotype 
male (1959. 1.20. 156) and one paratype female from pine and hardwood litter, 
St. Thomas, Portland Gap, 5,500 ft., 8 .iii. 1956, collected by P. F. Bellinger. The 
paratype has been returned to the collector. 



Sejus hulli sp. nov. 

Female. Dorsal shield (359 p long X 195 pi wide) completely covered by a 
conspicuous reticulated pattern (Text-fig. 143). "Anterior dorsal shield" with 
twenty-one pairs of simple setae, verticals on a prominent protuberance and directed 
anteriorly. " Posterior dorsal shield" with fifteen pairs of simple setae, J5 are 
short. All setae arise from small protuberances. The distribution and relative 
lengths of the dorsal setae and the ornamentation of the dorsal shield are shown in 
the Text-fig. 

Tritosternum well developed with pilose laciniae. Sternal shield with a very faint 



82 



THE PLATYSEIINAE (MESOSTIGMATA : ACEOSEJIDAE 



reticulated area medially, and bearing three pairs of simple setae (Text-fig. 144). 
Genital shield bearing a single pair of setae. Ventri-anal shield a little broader than 
long (94 fi long x 114 fi wide), reticulated and bearing nine simple setae. Between 
the genital and ventri-anal shields lie about six small sclerotized plates. Stigma 
situated between coxae III and IV ; no distinct post-stigmal prolongation of peri- 
treme ; peritrematal shield extending posterior to coxa IV. Metapodal plates 
small. 

Venter of gnathosoma with rostral and internal seta on palptrochanter long, 




Figs. 143-147. Sejus hulli sp. nov., female. Fig. 143, dorsum. Fig. 144, venter. 
Fig. 145, tectum. Fig. 146, chelicera. Fig. 147 tarsus II. 



whip-like. All setae on pedipalp simple. Tectum (Text-fig. 145) three-pronged. 
The dentition of the chelicera is shown in Text-fig. 146. 

Leg I (360 ju) with the tarsus (101 /i) nearly twice the length of the tibia (56 /i). 
Setae on tarsus and tibia fine, those on remaining segments stouter. Claws on leg I 
minute, tarsus swollen in its distal third. Tarsi II-IV with a pair of lanceolate 
setae, and ambulacra with acuminate lobes (Text-fig. 147). 

Male. Unknown. 

Locality. Jamaica, B.W.I. A single female (1959. 1.20. 157) collected in 
Lycopodium at Fairy Glade, St. Andrew, 6.XH.1956, by P. F. Bellinger. 



THE PLATYSEIINAE (MESOSTIGM AT A : ACEOSEJIDAE) 83 

Sejus jamaicensis sp. nov. 

Female. Dorsal shield (477 /i long x 310 jlc wide) concave posteriorly and 
covered with a light reticulated pattern (Text-fig. 148). " Anterior dorsal shield " 
with twenty-one pairs of simple setae ; verticals well separated, short, and arising 
from weak bases. " Posterior dorsal shield " with fifteen pairs of simple setae of 
which J5 are the shortest. The distribution of the dorsal setae and the ornamenta- 
tion of the dorsal shield are shown in the Text-fig. 




Figs. 148-152. Sejus jamaicensis sp. nov., female. Fig. 148, dorsum. Fig. 149, venter. 
Fig. 150 tectum. Fig. 151, chelicera. Fig. 152, tarsus II. 



Tritosternum well developed with pilose laciniae. Sternal shield with a little 
sculpturing and two circular areas anteriorly (Text-fig. 149) , and bearing three pairs 
of simple setae. Genital shield with one pair of simple setae. Ventri-anal shield 
wider than long (114-119 /i long X 139-144 /i wide), bearing nine simple setae, and 
only lightly sculptured. Between the genital and ventri-anal shields lie a row of 
four narrow platelets and, posterior to these, two smaller ones. Stigma situated 
between coxae III and IV ; peritreme extending posterior to coxa IV. Metapodal 
plates very small. 

Venter of gnathosoma with rostral and internal palptrochanter setae long, whip- 
like. All setae on pedipalp simple. Tectum (Text-fig. 150) three-pronged. The 
dentition of the chelicera is shown in Text-fig. 151. 

Leg I (924 fi long) with the tarsus (215 fi) only slightly shorter than the tibia 
(228 fi). Setae on tarsus, tibia and genu very fine ; those on remaining segments 



8 4 



THE PLATYSEIINAE (MESOSTIGMATA : ACEOSEJIDAE 



stouter. Claws minute, sessile. Tarsi II-IV with a pair of lanceolate setae ; 
ambulacra with three acuminate lobes (Text-fig. 152). 

Male. Unknown. 

Locality. Jamaica, B.W.I. The holotype female (1959. 1.20. 158) and one 
paratype female from wet leaves in spray zone, Cape Crow Falls, St. Andrew, 
31.iii.1956, collected by P. F. Bellinger. The paratype has been returned to the 
collector. 

Sejus neborealis sp. nov. 

Female. Dorsal shield (570 /i long x 330-340 fi wide) entirely covered by a 
strong reticulated pattern, and with a sclerotized tubercle in the region of setae 




Figs. 153-157. Sejus neborealis sp. nov., female. Fig. 153, dorsum. Fig. 154, venter. 
Fig. 155, tectum. Fig. 156, chelicera. Fig. 157, tarsus II. 

J3-J4 (Text-fig. 153). " Anterior dorsal shield " with twenty-one pairs of simple 
setae, vertical setae contiguous and directed anteriorly. "Posterior dorsal shield " 
with fifteen pairs of simple setae, J4 situated on the margin of the sclerotized tubercle. 
The distribution and relative lengths of the dorsal setae, and the ornamentation of 
the dorsal shield are shown in the Text-fig. 



THE PLATYSEIINAE (MESOSTIGM AT A : ACEOSEJIDAE) 85 

Tritosternum well developed with pilose laciniae. Sternal shield with a small 
median reticulated area anteriorly, and bearing three pairs of short simple setae 
(Text-fig. 154). Genital shield slightly convex posteriorly and bearing one pair of 
simple setae. Ventri-anal shield broader than long (172-177 /i long X 240-253 jll 
wide), wholly reticulated and bearing nine simple setae. Between the genital and 
ventri-anal shields lie four sclerotized platelets. Stigma situated between coxae III 
and IV with the posterior prolongation of the peritreme extending in line with the 
middle of coxa IV. Peritrematal shield well developed, extending posterior to coxa 
IV, and crenate externally. Metapodal plates small. 

Venter of gnathosoma with rostral and internal palptrochanter setae long, whip- 
like. All setae on pedipalp simple. Tectum (Text-fig. 155) three-pronged. The 
dentition of the chelicera is shown in Text-fig. 156. 

Leg I (734-774 fi long) with the tarsus (228-243 fi) approximately twice the length 
of the tibia (114-126 jlc). Some setae on tarsus I relatively long and standing at 
right angles to the segment ; ambulacrum short, claws small (and inconspicuous) ; 
setae of remaining segments stouter. Tarsi II-IV with a pair of lanceolate setae 
and ambulacrum with three acuminate lobes (Text-fig. 157). 

Male. Unknown. 

Locality. Jamaica, B.W.I. The holotype female (1959. 1.20. 158) and three 
paratype females (1959. 1.20. 159-160) collected in moss on concrete, Coolsbade, 
Portland, 17 . vii . 1956, by P. F. Bellinger. One paratype has been returned to the 
collector. 



Sejus nodosus sp. nov. 

Female. Dorsal shield (518-529 fi long x 310 fi wide) slightly attenuated in its 
posterior half, and covered by a reticulated pattern (Text-fig. 158). There is a 
pronounced sclerotized tubercle in the region of setae J3-J4. " Anterior dorsal 
shield " with twenty-one pairs of simple setae ; vertical setae closely situated and 
anteriorly directed. " Posterior dorsal shield " with twenty-one pairs of simple 
setae, of which J5 are the shortest. The distribution and relative lengths of the 
setae and the ornamentation of the dorsal shield are shown in the Text-fig. 

Tritosternum well developed and with pilose laciniae. Sternal shield plain and 
bearing three pairs of simple setae (Text-fig. 159). Genital shield slightly convex 
posteriorly, reticulated, and bearing a single pair of setae. Ventri-anal shield 
wider than long (114-116 ju, long x 134-144 fi wide), lightly sculptured and bearing 
nine simple setae. Between the genital and ventri-anal shields lies a row of two to 
four narrow sclerotized platelets. Stigma situated between coxae III and IV ; 
peritreme not extending posterior to the stigma. Metapodal plates small. 

Venter of gnathosoma with rostral and internal palptrochanter setae long, whip- 
like. All setae on pedipalp simple. Tectum (Text-fig. 160) three-pronged, and 
chelicerae typical of the genus. 

Leg I (c. 620 fi long) with the tarsus (165 fi) longer than the tibia (126 fi). Tarsus 
I with a strong erect spine dorsally, remaining setae fine ; tarsal claws small but 



86 



THE PLATYSEIINAE (MESOSTIGMATA : ACEOSEJIDAE) 



conspicuous. Tarsi II— III with a pair of acuminate lobes (Text-fig. 161.) Leg IV is 
about 913 /i long, with the tarsus 392 jlc long, markedly elongate, and three times as 
long as the tibia. 
Male. Unknown. 




Figs. 



1 58-1 61. Sejus nodosus sp. nov., female. Fig. 158, dorsum. Fig. 159, venter. 
Fig. 160, tectum. Fig. 161, tarsus II. 



Locality. Jamaica, B.W.I. The holotype female (1959. 1.20. 161) and one 
paratype from moss on concrete, Coolsbade, Portland, 17.vii.1955, collected by 
P. F. Bellinger. The paratype has been returned to the collector. 



Sejus browningi sp. nov. 

Female. Dorsal shield (330-354 jlc long x 190-210 fi wide) very heavily reticu- 
lated (Text-fig. 162). "Anterior dorsal shield" with twenty-one pairs of setae; 
"posterior dorsal shield" with fifteen pairs of setae. Dorsal setae J5 are short 
and simple, but the remainder are lanceolate (Text-fig. 163). The highest part of 
the dorsal shield is in the region of setae J1-J3. The vertex is produced anteriorly. 
The distribution and relative lengths of the dorsal setae are shown in Text-fig. 162. 



THE PLATYSEIINAE (MESOSTIGM AT A : ACEOSEJIDAE) 



87 



Tritosternum with a pair of pilose laciniae. Sternal shield with a narrow longi- 
tudinal reticulated area anteriorly (Text-fig. 164) and bearing three pairs of simple 
setae. Genital shield broad, truncated posteriorly, lightly striated and bearing a 
single pair of simple setae. Ventri-anal shield wider than long (99-101 /i long 
X 121-139 fi wide), lightly reticulated anteriorly, and bearing nine simple setae. A 
row of five narrow sclerotized plates lies on the membrane between the genital and 
ventri-anal shields. Stigma situated between coxae III and IV, with the peritreme 




Figs. 162-167. 
dorsal seta. 
tarsus II. 



Sejus browningi sp. nov., female. Fig. 162, dorsal shield. Fig. 163, 
Fig. 164, venter. Fig. 165, tectum. Fig. 166, chelicera. Fig. 167, 



extending only a short distance posterior to the stigma. Peritrematal shield 
extending posterior to coxa IV. Metapodal shields triangular and situated close to 
the peritrematal shield. 

Venter of gnathosoma with rostral and internal palptrochanter setae long, whip- 
like. All setae on pedipalp simple. Tectum (Text-fig. 165) three-pronged and 
typical of the genus. The dentition of the chelicera is shown in Text-fig. 166. 

Leg I (320-340 fi long) with the tarsus (88 /i) longer than the tibia (58 ja). Setae 
on tarsus I very fine ; those on remaining segments somewhat stouter. Tarsus I 
is swollen in its distal third ; without ambulacrum. Tarsi II-IV with a pair of 
lanceolate setae, and ambulacra with lateral lobes acuminate ; median lobe strongly 
produced and rounded apically (Text-fig. 167). 



88 



THE PLATYSEIINAE (MESOSTIGMAT A : ACEOSEJIDAE) 



Male. Unknown. 

Locality. Sikkim and Argentina. The holotype female (1959. 1.20. 162) and 
two paratype females (1959. 1.20. 163-164) from leaves and leaf-mould in a wood 
on steep hillside, Chungtang, Sikkim, 5,120 ft., 16. ii. 1952, collected by Dr. Theresa 
Clay ; and another paratype female (1959 . 1 . 20 . 165) from rotting vegetable matter, 
Tucuman City, Argentina, 500 m., January, 1953, collected by Dr. P. Wygodzynsky. 



Sejus ornatus sp. nov. 

Female. Dorsal shield (395-400 /i long x 245-253 ju, wide) with lateral incisions 
between setae S2 and S3, and covered by a network of small tubercles (Text-fig. 
168). Highest part of dorsal shield in the region of setae J1-J3. " Anterior dorsal 
shield " with twenty- two pairs of simple setae ; " posterior dorsal shield " with fifteen 
pairs of simple setae. Setae J5 are long. Vertex of dorsal shield produced ante- 




Figs. 168-173. Sejus ornatus sp. nov., female. Fig. 168, 
Fig. 170, vertex. Fig. 171, tectum. Fig. 172, chelicer 



dorsum. Fig. 169, venter. 



Fig. 172, chelicera. Fig. 173, tarsus II. 



THE PLATYSEIINAE ' (MESOSTI GM AT A : ACEOSEJIDAE) 89 

riorly. The distribution and relative lengths of the dorsal setae and the ornamenta- 
tion of the dorsal shield are shown in the Text-fig. 

Tritosternum well developed, with pilose laciniae. Sternal shield with a circular 
impression antero-medially and bearing three pairs of simple setae (Text-fig. 169). 
Genital shield broad, truncated posteriorly, and bearing a single pair of setae. 
Ventri-anal shield wider than long (1 18-139 fi long X 164-177 ju, wide), ornamented 
with small tubercles and punctations and bearing nine simple setae. Stigma 
situated between coxae III and IV ; peritremes not joined anteriorly (Text-fig. 170), 
and not extending posterior to the stigmata. Outer margin of peritreme crenate ; 
peritrematal shield extending to the posterior level of coxa IV. Metapodal shields 
narrow and transversely situated. 

Venter of gnathosoma with rostral and internal palptrochanter setae long, whip- 
like. All setae on pedipalp simple. Tectum (Text-fig. 171) three-pronged, typical 
of the genus. The dentition of the chelicera is shown in Text-fig. 172. 

Leg I (360-366 ju long) with the tarsus (103-113 /i) longer than the tibia (58-63 jlc). 
Setae on tarsus I very fine ; those on remaining segments stouter. Tarsi II-IV 
with a pair of lanceolate setae, and ambulacra with lateral lobes acuminate, median 
lobe rudimentary (Text-fig. 173). Tarsus I slightly swollen at its distal half, 
pul villus and claws small. 

Male. Unknown. 

Locality. Argentina. The holotype female (1959. 1.20. 166) and three para- 
types (1959. 1.20. 167-169) from horse dung and decomposing vegetable matter in 
forest, Quebrada de los Sosa, Tafi, Tucuman, 1,300 m., 17. v. 1953, collected by 
Dr. P. Wygodzynsky. 



Sejus tuberculatus sp. nov. 

Female. Dorsal shield (477-518 /i long x 310-330 /i wide) minutely punctured 
and slightly concave posteriorly (Text-fig. 174). "Anterior dorsal shield" with 
twenty-one pairs of simple setae. " Posterior dorsal shield " with fifteen pairs of 
simple setae ; J5 are the shortest. The distribution and relative lengths of the 
setae, and the ornamentation of the dorsal shield are shown in the Text-fig. 

Tritosternum well developed and with pilose laciniae. Sternal shield with faint 
sculpturing, and bearing three pairs of simple setae (Text-fig. 175). Genital shield 
flask-shaped, and bearing a single pair of setae. Ventri-anal shield almost circular 
in outline (130-144 /i long x 144-147 jli wide) and bearing nine simple setae. Between 
the genital and ventri-anal shields lies a transverse row of about three narrow plates, 
and posterior to these lie two small plates. Stigma situated between coxae III and 
IV ; peritreme and peritrematal shields extending posterior to coxa IV. Metapodal 
plates small. 

Venter of gnathosoma with rostral and internal palptrochanter setae long, whip- 
like. All setae on pedipalp simple. Tectum (Text-fig. 176) three-pronged. The 
dentition of the chelicera is shown in Text-fig. 177. 

Leg I (794-804 jlc long) with the tarsus (162-177 ji) shorter than the tibia (182- 

ZOOL. 6, NO. 2. 7 



go 



THE PLATYSEIINAE (MESOSTIGMATA : ACEOSEJIDAE) 



197 fi). Setae on tarsus, tibia and genu fine ; those on other segments stouter 
tarsal claws minute, inconspicuous. Tarsi II-IV with a pair of lanceolate setae 
ambulacra with three acuminate lobes (Text-fig. 178). 
Male. Unknown. 




I/O 

Figs. 174-178. Sejus tuberculatus sp. nov., female. Fig. 174, dorsum. Fig. 175, venter 
Fig. 176, tectum. Fig. 177, chelicera. Fig. 178, tarsus II. 



Locality. Argentina. The holotype female (1959 . 1 . 20 . 170) and two paratype 
females (1959. 1.20. 171. 172) from decaying vegetable matter in forest, Quebrada 
de los Sosa, Tafi, Tucuman, 13. v. 1953, 1,300 m., collected by Dr. P. Wygodzynsky. 

The following species also appear to belong to the genus Sejus, but with the 
exception of Episeius groenlandicus, have not been examined by us : 

Acarus tendens Schrank, 1803. Fauna Boica, 3 : 209. Oudemans (1929) con- 
siders this species to be synonymous with Platyseius subglaber (Ouds.). 

Gamasus scabriculus Nordmann, 1832. Mikr. Beitr. Naturg. wirbett. Thiere : 85. 

Epicrius glaber Berlese, 1886. A. M.S., 30, No. 9. Italy. 

Hypoaspis scutulis Banks, 1914. Psyche, 21 : 161. Brazil. 

Lasioseius borealis var. temperatus Berlese, 1916. Redia, 12 : 34. Palermo. 

Lasioseius grandis Berlese, 1916. Tom. cit. : 34. Piemonte, Venice. 

Lasioseius parapodicus Berlese, 1916. Tom. cit. : 35. Java. 

Lasioseius similis Berlese, 1916. Tom. cit. : 35. Italy. This species may be a 



THE PLATYSEIINAE (MESOSTIGM AT A : ACEOSEJIDAE) 91 

synonym of Sejus serratus (Halbert). 

Lasioseius mutilus Berlese, 1916. Tom. cit. : 37. Florence. 

Lasioseius (Cheiroseius) alpestris Berlese, 1916. Tom. cit. : 41. Italy. 

Episeius major incisus Willmann, 1938. Ann. Hist. nat. Mus. Hung., 31 : 168. 
Hungary. 

Episeius groenlandicus Haarlov, 1942. Medd. Grenland, 128 : 18. Greenland. 

Episeius aequalis Schweizer, 1949. Res. Rech. sci. Pare Nat. Suisse, N.F. 2 : 64. 
Switzerland. 

Episeius handschini Schweizer, 1949. Tom. cit. : 71. Switzerland. 

Episeius salicorniae Willmann, 1949. Veroff. Mus. Nat. Bremen, No. iA : 118. 
Poland. 

Episeius longipes Willmann, 1951. SB. ost. Akad. Wiss., Abt. 1, 160 : 112. 
Austria. 

Platyseius mackerassae Womersley, 1956. J. Linn. Soc. Lond. (Zool.) 42 : 552. 
Queensland. 

Genus ZERCONOPSIS HuU 

Zerconopsis Hull, J. E., 1918, Trans, nat. Hist. Soc. Northumb., 5 : 65. 

Platyseiine mites with the "posterior dorsal shield" bearing fourteen pairs of 
setae comprising five pairs in the J and the Z series but only four pairs in the S 
series (S2 being situated on the lateral interscutal membrane). Three to five pairs 
of dorsal setae stout and paddle-like (Text-figs. 179, 192). Lateral margins of the 
shield entire or incised anterior to Si. Sternal shield in the female with three pairs 
of setae ; metasternals situated on platelets. Genital shield wedge shaped, genital 
setae situated on or off the shield. Ventri-anal shield large and bearing from three 
to seven pairs of setae in addition to the three setae normally associated with the 
anus. Male with sterniti-genital and free ventri-anal shield. Peritreme without 
post-stigmatic process ; peritrematal shield poorly developed posterior to coxa IV. 
Chaetotaxy of the venter of the gnathosoma and of the pedipalps typical of the 
subfamily. Chelicerae chelate-dentate, movable digit in the male with a short 
spermatophoral process ; tectum basically three-pronged. Legs I-IV with a well- 
developed ambulacrum ; median lobes of pulvilli rounded apically. 



Type species Gamasus remiger Kramer, 1876 
Key to Species — Adults 

1. Dorsal shield with distinct lateral incisions anterior to Si (Text-figs. 184 and 197) 2 
— Lateral margins of the dorsal shield entire ....... 3 

2. Genital shield in the female narrow, without setae (Text-fig. 198). Labrador 

Z.l abradorensis sp. n. (p. 99) 
— Genital shield in the female broad, with a pair of setae (Text-fig. 185). Switzerland 

Z. muestairi (Schweizer) (p. 94) 

3. Dorsal shield with five pairs of paddle-like setae (Text-fig. 192) ; genital shield 
without setae (Text-fig. 193). Hungary . . Z. decemremiger sp. n. (p. 96) 



92 THE PLATYSEIINAE (MESOSTIGM AT A : ACEOSEJIDAE) 

— Dorsal shield with three pairs of paddle-like setae (Text-figs. 179 and 189) ; genital 

shield with or without setae .......... 4 

4. Genital shield with a pair of setae (Text-fig. 180) ; with one pair of large, subtriangular 

metapodal shields. Europe ..... Z. remiger (Kramer) (p. 92) 

— Genital shield narrow, without setae (Text-fig. 190) ; with two pairs of metapodal 

shields. England ....... Z. michaeli sp. n. (p. 95) 



Zerconopsis remiger (Kramer) 

Gamasus regimer Kramer, P., 1876, Arch. Naturg., 42 : 93. 
Ameroseius bispinosus Berlese, A., 1910, Redia, 6 : 253. 

Female. Dorsal shield (570-630 /i long X 310-382 jlc wide) covered antero- 
laterally with a distinct reticulated pattern ; posterior and central areas with a 
lighter network (Text-fig. 179). " Anterior dorsal shield " with twenty-two pairs of 
setae of which one pair is paddle-like, the remainder simple ; verticals short, each 
on a tubercle. " Posterior dorsal shield " with fourteen pairs of setae of which two 
pairs are paddle-like and the remainder simple ; J5 are the shortest. The distribu- 
tion and relative lengths of the dorsal setae and the ornamentation of the dorsal 
shield are shown in Text-fig. 179. 

Tritosternum with a narrow base and pilose laciniae, and flanked by a narrow 
pair of pre-endopodal shields. Sternal shield with a fine granular pattern and 
bearing three pairs of simple setae (Text-fig. 180). Genital shield also with granular 
markings, and bearing one pair of setae. Ventri-anal shield wider than long (220- 
223 fi long x 225-303 /i wide), heavily reticulated and bearing thirteen simple setae. 
Between the genital and ventri-anal shields lie six platelets. Stigma situated 
between coxae III and IV with the peritreme not extending posterior to the stigma. 
Metapodal plates conspicuous. 

Venter of gnathosoma with rostral and internal palptrochanter setae long, whip- 
like. All setae on pedipalp simple. Tectum three-pronged and typical of the 
genus. The dentition of the chelicera is shown in Text-fig. 181. 

Leg I (379-400 fi long) with the tarsus (99-103 /i) longer than the tibia (63-68 jlc) ; 
claws small, pul villus short. Tarsus I smooth with fine setae ; tibia, genu and 
femur with majority of setae arising from strong tubercles. Tarsi II-IV with a 
pair of lanceolate setae ; ambulacra with median lobes of pulvilli rounded apically. 

Male. Dorsal shield (530-550 /i long x 299-310 /i wide) with chaetotaxy and 
ornamentation similar to the female. Sterniti-genital shield with five pairs of simple 
setae. Ventri-anal shield (Text-fig. 182) reticulated anteriorly and bearing seventeen 
simple setae. The gnathosoma, tectum, pedipalps and legs are similar to those of 
the female. Leg I (c. 420 fi long) with the tarsus (99 p) and the tibia (68 /i). Sperma- 
tophoral process about a third as long again as the movable digit of the chelicera 
(Text-fig. 183). 

Distribution. Kramer (1876) described this species from under fallen leaves 
without giving a locality, and Berlese (1910) described Ameroseius bispinosus from 
rotting leaves at Palermo. We have examined specimens from Nadap, Herkules- 



THE PLATYSEIINAE (MESOSTIGM AT A : ACEOSEJIDAE 



93 



furdo, and Gyertyanliget in Hungary, and British specimens from Goyt Valley, 
Cheshire, 24. xi. 1940, collected by H. Britten; Rydal Water, Westmorland, 
29. xi. 1954, collected by M. E. Bacchus ; Harefield, Middlesex, 22. i. 1956, collected 
by A. H. G. Alston; and Avonmouth, Gloucestershire, n. v. 1957, collected by 
P. N. Lawrence. 




Figs. 179-183. Zerconopsis remiger (Kramer). Fig. 179, dorsum of female. Fig. 180, 
venter of female. Fig. 181, chelicera of female. Fig. 182, ventri-anal shield of male. 
Fig. 183, chelicera of male. 



94 



THE PLATYSEIINAE (MESOSTIGM ATA : ACEOSEJIDAE; 



Zerconopsis muestairi (Schweizer) comb. nov. 

Lasioseius mustairi Schweizer, J., 1949, Rec. Rech. sci. Pare Nat. Suisse N.F. 2 : 50. 

Female. Dorsal shield (585-630 /i long x 315-378 fi wide) with a lateral incision 
anterior to seta Si ; lightly reticulated anteriorly and entirely granular (Text-fig. 
184). " Anterior dorsal shield " with nineteen pairs of setae of which one pair is 
paddle-like, remainder simple ; verticals relatively long and well separated. " Pos- 
terior dorsal shield " with fourteen pairs of setae of which two pairs are paddle-like, 




Figs. 184-188. Zerconopsis muestairi (Schweizer). Fig. 184, dorsum of female. Fig. 
185, venter of female. Fig. 186, tectum of female. Fig. 187. ventri-anal shield of male. 
1 [ 1 £8, chelicera of male. 



THE PLATYSEIINAE (MESOSTIGM ATA : ACEOSEJIDAE) 95 

the remainder simple ; J5 are the shortest. The distribution and relative lengths 
of the dorsal setae and the form of the dorsal shield are shown in the Text-fig. 

Tritosternum with a narrow base and pilose laciniae. Sternal shield with faint 
sculpturing and bearing three pairs of simple setae (Text-fig. 185). Genital shield 
also with faint markings, and bearing a single pair of setae. Ventri-anal shield 
wider than long (180-223 /* l° n g X 240-270 fi wide), reticulated in its anterior half 
and bearing thirteen simple setae. Between the genital and ventri-anal shields lie 
six platelets. Stigma situated between coxae III and IV with the peritreme not 
extending posterior to the stigma. Metapodal plates small. 

Venter of gnathosoma with rostral and internal palptrochanter setae long, whip- 
like. All setae on pedipalp simple. Tectum (Text-fig. 186) three-pronged and 
typical of the genus. The chelicerae are figured by Schweizer (1949). 

Leg I (432-504 fi long) with the tarsus (1 18-132 ju) longer than the tibia (68-75 /i). 
Claws on leg I distinct, pulvillus small ; tarsal setae fine, those on other segments 
stouter. Tarsi II-IV with a pair of lanceolate setae ; median lobes of pulvilli 
rounded apically. 

Male. Dorsal shield (467-498 /i long X 252-289 fi wide) with chaetotaxy and 
ornamentation similar to the female. Sterniti-genital shield with five pairs of 
simple setae. Ventri-anal shield with nineteen simple setae (Text-fig. 187). The 
gnathosoma, tectum, pedipalps and legs are typical. Leg I (326-332 fi long) with 
the tarsus 108-116 fi and the tibia 63-65 /i. The spermatophoral process and 
chelicera are shown in Text-fig. 188. 

Locality. This species is known only from the Swiss National Park. We have 
examined five females and three males from the type series. 



Zerconopsis michaeli sp. nov. 

Female. Dorsal shield (498 /i long x 253 /i wide) with a heavy distinct pattern 
of depressions ; lateral margins irregular, posterior margin crenate (Text-fig. 189). 
" Anterior dorsal shield " with nineteen pairs of setae of which one pair is paddle- 
like, the remainder simple ; verticals on tubercles and directed anteriorly. " Pos- 
terior dorsal shield " with fourteen pairs of setae of which two pairs are paddle-like, 
the remainder simple. The distribution of the setae and the structure and orna- 
mentation of the dorsal shield are shown in the Text-fig. 

Tritosternum with a narrow base and pilose laciniae. Sternal shield plain, 
bearing three pairs of simple setae (Text-fig. 190). Genital shield narrow, setae off 
the shield. Ventri-anal shield wider than long (160 ji long x 212 /i wide), with a 
pattern of reticulations and dots, and bearing eleven setae of which the post-anal is 
long and stout. There are at least two narrow sclerotized platelets between the 
genital and ventri-anal shields. Stigma situated between coxae III and IV ; no 
post-stigmal prolongation of the peritreme. Metapodal plates small. 

Venter of gnathosoma with rostral and internal palptrochanter setae long, whip- 
like. All setae on pedipalp simple. Tectum (Text-fig. 191) three-pronged and 
typical of the genus. Chelicerae typical. 



9 6 



THE PLATYSEIINAE (MESOSTI GM AT A : ACEOSEJIDAE) 



Leg I (387 11 long) with the tarsus (101 /i) almost twice the length of the tibia 
(58 jj). Legs II-IV incomplete in the unique specimen, but some of the setae arise 
from tubercles. 

Male. Unknown. 

Locality. A single female (1930 . 8 . 25 . 1840) from "England" in the A. D. 
Michael Collection and labelled " Sejus remiger Kramer ". 




Figs. 189-191. Zerconopsis michaeli sp. nov., female. Fig. 189, dorsal shield. Fig. 190, 

venter. Fig. 191, tectum. 



Zerconopsis decemremiger sp. nov. 

Female. Dorsal shield (590-600 jlc long x 360-382 ja wide) with a dense pattern 
of depressions, largest around the anterior margin and becoming smaller posteriorly 
(Text-fig. 192). " Anterior dorsal shield " with twenty pairs of setae of which two 
pairs are paddle-like, the remainder simple ; verticals separated by about three times 
the diameter of their bases. " Posterior dorsal shield " with fourteen pairs of setae 
of which three pairs are paddle-like ; J5 are the shortest. The distribution of the 
setae and the ornamentation of the dorsal shield are shown in the Text-fig. 

Tritosternum with a narrow base and pilose laciniae. Sternal shield with faint 



THE PLATYSEIINAE (MESOSTIGM AT A : ACEOSEJIDAE) 97 




Figs. 192-196. Zerconopsis decemremiger sp. nov. Fig. 192, dorsal shield of female. 
Fig. 193, venter of female. Fig. 194, tectum of female. Fig. 195, ventri-anal shield of 
male. Fig. 196, chelicera of male. 



98 THE PLATYSEIINAE (MESOSTIGMATA : ACEOSEJIDAE) 

reticulations and bearing three pairs of simple setae (Text-fig. 193). Genital shield 
reticulated ; genital setae off the shield. Ventri-anal shield wider than long (205- 
220 p long X 253-263 /i wide), with a conspicuous pattern of small depressions and 
bearing eleven simple setae. Between the genital and ventri-anal shields lie four 
small plates. Stigma situated between coxae III and IV ; peritreme without 
post-stigmal prolongation. Metapodal plates small. 

Venter of gnathosoma with rostral and internal palptrochanter setae long, whip- 
like. All setae on pedipalp simple. Tectum three-pronged and typical of Zerconop- 
sis (Text-fig. 194) : chelicerae also typical. 

Leg I (434 p long) with the tarsus (111-116 p) longer than the tibia (68^70 p). 
Claws on leg I small ; tarsal setae fine, those on remaining segments somewhat 
stouter and arising mainly from small tubercles. Tarsi II-IV with a pair of lanceo- 
late setae. Ambulacra with median lobes of pul villi rounded apically. 

Male. With dorsal shield (507-550 p long x 289-330 p wide) essentially the 
same as the female. Sterniti-genital shield with five pairs of simple setae. Ventri- 
anal shield (Text-fig. 195) similar in ornamentation to the female and bearing 
nineteen simple setae. The gnathosoma, tectum, pedipalps and legs are similar to 
those of the female. Leg I (434 p long) with the tarsus (111-113 p) longer than the 
tibia (68-70 p). The spermatophoral process and chelicera are shown in Text-fig. 
196. 

Locality. Hungary. The holotype female (1959. 1.20. 177), allotype male 
(1959. 1. 20. 178), four female and two male paratypes (1959. 1.20. 179-184) from 
Herkulesfurdo, April- June, 1938 ; and four female and two male paratypes from 
Gyertyanliget, 6-i7th August, 1940, sent to us by Dr. J. Balogh, Budapest. 



Zerconopsis labradorensis sp. nov. 

Female. Dorsal shield (581-609 p long x 299-320 p wide) with a lateral incision 
anterior to seta Si, reticulated antero-laterally and the remainder granulate (Text- 
fig. 197). " Anterior dorsal shield" with twenty-one pairs of setae of which one 
pair is paddle-like, the remainder simple ; vertical setae with their bases two 
diameters apart. " Posterior dorsal shield " with fourteen pairs of setae of which 
two pairs are paddle-like ; J5 are the shortest. The distribution of the setae and 
the form of the dorsal shield are shown in the Text-fig. 

Tritosternum with a narrow base and a pair of pilose laciniae. Sternal shield 
almost without markings, and bearing three pairs of simple setae (Text-fig. 198). 
Genital shield plain ; setae off the shield. Ventri-anal shield wider than long (177- 
215 p long x 253-260 p wide), reticulated anteriorly and granulate posteriorly, and 
bearing fifteen simple setae. Between the genital and ventri-anal shields lie six 
small sclerotized plates. Stigma between coxae III and IV ; peritreme without 
post-stigmal prolongation (Text-fig. 199). Metapodal plates small. 

Venter of gnathosoma with rostral and internal palptrochanter setae long, whip- 
like. All setae on pedipalp simple. Tectum (Text-fig. 200) three-pronged ; cheli- 
cerae typical (Text-fig. 201). 



THE PLATYSEIINAE (MESOSTIGM ATA : ACEOSEJIDAE) 99 




Figs. 197-204. Zerconopsis labradorensis sp. nov. Fig. 197, dorsum of female. Fig. 
198, venter of female. Fig. 199, peritrematal shield of female. Fig. 200, tectum of 
female. Fig. 201, chelicera of female. Fig. 202, tarsus II of female. Fig. 203, venter 
of male. Fig. 204, chelicera of male. 



ioo THE PLATYSEIINAE (MESOSTIGM AT A : ACEOSEJIDAE) 

Leg I (462-502 /i long) with the tarsus (124-134 ju) twice the length of the tibia 
(65-73 ji) ; tarsal claws small ; setae fine, those on remaining segments shorter and 
stouter. Tarsi II-IV (Text-fig. 202) with a pair of lanceolate setae ; ambulacra 
with median lobes of pulvilli rounded apically. 

Male. Dorsal shield (488 ju, long x 238 jlc wide) essentially the same as in the 
female. Sterniti-genital shield with five pairs of simple setae. Ventri-anal shield 
(Text-fig 203) similar in ornamentation to the female and bearing nineteen simple 
setae. The gnathosoma, tectum, pedipalps and legs are similar to those of the 
female. Leg I (462 /i long) with the tarsus (124 fi) twice as long as the tibia (65 ju). 
The spermatophoral process of the chelicera is shown in Text-fig. 204. 

Locality. Labrador. The holotype female (1959. 1.20. 185), allotype male 
(1959. 1. 20. 186) and six paratype females (1959. 1.20. 187-192) from wet willow 
humus, West Carter Basin, 4-17. viii. 1958, collected by P. N. Lawrence during the 
British Schools' Exploring Society Expedition. 



SUMMARY 

1. This paper deals with the external morphology and classification of the Platy- 
seiinae. Four genera, namely, Sejus C. L. Koch (syn. Cheiroseius, Episeius and 
Episeiella), Platyseius Berlese, Zerconopsis Hull, and Plesiosejus gen. no v. are 
recognized. Keys are given to the species represented in the Collections of the 
British Museum (Natural History). 

2. The following new synonymy is introduced : 

Lasioseius (Platyseius) capillatus Berl. = Hypoaspis subglabra Oudms. 

Paraseius tenuipes Halbert = Ameroseius italicus Berl. 

Episeiella heteropoda Willm. = Sejus viduus C. L. Koch 

Episeius montanus Willm. = Ameroseius borealis Berl. 

Episeius ovaspini Schweizer = Lasioseius (Episeius) glaber var. curtipes Halbt. 

Parasejus glaber var. minor Tragardh = Epicrius laelaptoides Berl. 

Lasioseius (Episeius) sphagni Halbt. = Epicrius laelaptoides Berl. 

3. The following twenty-seven new species are described and figured : Platyseius 
jamaicensis ; Platyseius spinosus ; Plesiosejus horridus ; Sejus aciculatus ; S. 
alstoni ; S. antillanus ; 5. areolatus ; S. bellingeri ; 5. browningi ; S. cassiteridum ; 
S. clayi ; S. dromadis ; S. hulli ; S. jamaicensis ; S. kennedyi ; S. neborealis ; 
S. nepalensis ; S. nodosus ; S. ornatus ; S. parbatensis ; S. phalangioides ; 5. 
signatus ; S. spinipes ; 5. tuberculatus ; Zerconopsis decemremiger ; Z. labradorensis 
and Z. michaeli. 

REFERENCES 

Berlese, A. 1882-1903. Acari, myriapoda et scorpiones hucusque in Italia reperta. Portici 
et Padua. 

1904. Acari nuovi II. Redia, 1 : 258-280. 

1905. Acari nuovi V. Ibid. 2 : 231-238. 

1910. Acari nuovi VI. Ibid. 6 : tav. 19, figs. 35, 35a. 

1910. Lista di nuove specie e nuovi generi di Acari. Ibid. 6 : 243-271. 



THE PLATYSEIINAE (MESOSTI GM AT A : ACEOSEJIDAE) 101 

BERLESE, A. 1913. Acari nuovi VII-VIII. Ibid. 9 : 77-1 11. 

1916. Centuria prima di Acari nuovi. Ibid. 12 : 19-67. 

Buitendijk, A. M. 1945. Voorloopige Catalogus van de Acari in de Collectie-Oudemans. 

Zool. Med. 23 : 281-391. 
Evans, G. O. 1957. An introduction to the British Mesostigmata (Acarina) with keys to 

families and genera. /. Linn. Soc. Lond. [Zool.) 43 : 203-259. 
1958. A revision of the British Aceosejinae (Acarina : Mesostigmata). Proc. zool. Soc. 

Lond. 131 : 177-229. 
Franz, H. 1943. Die Landtierwelt der Mittleren Hohen Tauern. Denkschr. Akad. Wiss. 

Wien, 107 : 79-119. 
Halbert, J. N. 191 5. Clare Island Survey Part 39U Acarinida ii — Terrestrial and marine 

Acarina. Proc. R. Irish Acad. 31 : 45-136. 
1923. Notes on Acari with descriptions of new species. /. Linn. Soc. Lond. (Zool.) 35 : 

363-392. 
Hirschmann, W. 1957- Acarologie — Gangsystematik der Parasitiformes Teil i. Rumpf- 

behaarung und Ruckenfldchen. Schriftenreihe fur vergleichende Milbenkunde. Furth/ 

Bay : 1-20 i-v, pis. 1-26. 
Hull, J. E. 1918. Terrestrial acari of the Tyne Province. Trans, nat. Hist. Soc. Northumb. 

[n.s.) 5 : 13-88. 
Koch, C. L. 1836-1844. Deutschlands Crustaceen, Myriapoden und Arachniden. Regens- 

burg. 

1837-1850. Obersicht des Arachnidensy stems. Nurnberg. 

Kramer, P. 1876. Zur Naturgeschichte einiger Gattungen aus der Familie der Gamasiden. 

Arch. Naturg. 42 : 46-105. 
Oudemans, A. C. 1902. Acariden von Borkum und Wangeroog. Abh. nat. Ver. Bremen, 

18 : 77-98. 

1903. Acarologische Aanteekeningen, VI. Ent. Ber. 1 : 83-88. 

1929. Kritisch Historisch Overzicht der Acarologie, Leiden, 2 : 87 

1936. Ibid 3A : 1-430. 

Schweizer, J. 1922. Beitrag zur Kenntnis der terrestrischen Milbenfauna der Schweiz. 

Verh. Naturf. Ges. Basel, 33 : 23-112. 
1924. Beitrage zur Kenntnis der Tierwelt norddeutscher Quellgebiete. Acarina (land- 

milben). Arch, fur Hydrobiol. 15 : 125-132. 
1949. Die Landmilben des Schweizerischen Nationalparkes. 1. Parasitiformes. Res. 

Rech. sci. Pare Nat. Suisse, N.F. 2 : 1-99. 
Sellnick, M. 1944. Zercon C. L. Koch. A cari-Bldtter fur Milbenkunde. Konigsberg, No. 5 : 

30-41. 
Tragardh, I. 1910. Acariden aus dem Sarekgebirge. Naturw. Untersuch. Sarekgeb. 4 : 375- 

586. 
1946. Outlines of a new classification of the Mesostigmata (Acarina) based on comparative 

morphological data. Acta Univ. Lund, N.F. Avd. 2, 42, 4 : 1-32. 
Vitzthum, H. G. 1 93 1. Terristrische Acarinen (unter Ausschluss der Oribatiden und Ixodiden) 

der Deutschen Limnologischen Sunda-Expedition. Arch, fur Hydrobiol., Suppl., 9 : 59- 

134- 

Willmann, C. 1938. Beitrag zur Kenntnis der Acarofauna des Komitates Bars. Ann. Hist. 

nat. Mus. Hung. 31 : 144-172. 
1949. Beitrage zur Kenntnis des Salzgebietes von Ciechocinek. Verojf. Mus. Nat. 

Bremen, No. \A : 106-135. 
1952. Die Milbenfauna der Nordsee-Insel Wangerooge. Veroff. Inst. Meeresf. Bremer- 

haven, 1 : 139-186. 
1953- Neue Milben aus den ostlichen Alpen. SB. ost. Akad. Wiss. 162 : 449-519. 



2 3 FEB I960 



BRITISH SLUGS 

(PULMONATA ; TESTACELLIDAE : 
ARIONIDAE, LIMACIDAE) 



H. E. QUICK 




BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY Vol. 6 No. 3 

LONDON: i960 




BRITISH SLUGS 

PULMONATA ; TESTACELLIDAE : 
ARIONIDAE, LIMACIDAE) 



BY 

H. E. QUICK 




Pp. 103-226 ; 2 Plates ; 19 Text-figures ; 23 Maps 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ZOOLOGY Vol. 6 No. 3 

LONDON: i960 



THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted in 1949, is 
issued in five series, corresponding to the Departments 
of the Museum, and an Historical series. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

This paper is Vol. 6, No. 3 of the Zoological series. 



Trustees of the British Museum, i960 



PRINTED BY ORDER OF THE TRUSTEES OF 
THE BRITISH MUSEUM 

Issued March, i960 Price Forty-five Shillings 



BRITISH SLUGS 

(PULMONATA ; TESTACELLIDAE, 

ARIONIDAE, LIMACIDAE 



By H. E. QUICK 





CONTENTS 










Page 




Introduction ........ . . 106 




Arrangement 










107 




Acknowledgments . 










107 




Native and Naturalized Species 










108 




Order Stylommatophora . 










108 




Superfamily Oleacinacea 










108 




Family Testacellidae 










108 




TESTACELLA Draparnaud 






108 




Testacella maugei Ferussac 






no 




Testacella haliotidea Draparnaud 






i*5 




Testacella sciitulum Sowerby 






117 




Superfamily Endodontacea .... 






119 




Family Arionidae ..... 






119 




Subfamily Arioninae .... 






119 




GEO MA LA C US Allman 






119 




Geomalacus maculosus Allman 






120 




ARION Ferussac 






124 




Avion intermedins Normand . 






125 




Avion fasciatus (Nilsson) 






127 




Avion hovtensis Ferussac 






130 




Avion subfuscus (Draparnaud) 






*33 




Avion lusitanicus Mabille 






135 




Avion atev atev (Linnaeus) 






138 




Avion atev vufus (Linnaeus) . 






H5 




Superfamily Zonitacea ..... 






147 




Family Limacidae ..... 






147 




Subfamily Parmacellinae 






149 




MILAX Gray .... 






149 




Milax gagates (Draparnaud) . 






149 




Milax cf. insulavis (Lessona & Pollonera] 






156 




Milax sowevbyi (Ferussac) 






158 




Milax budapestensis (Hazay) 






160 


ZOOL 


• 6, 3. 













io6 



BRITISH SLUGS 

















Page 


Subfamily Limacinae . . . . . .163 


A GRIOLIMA X Morch . 








163 


Agriolimax reticulatus (Miiller) 








164 


Agriolimax agvestis (Linnaeus) 








170 


Agriolimax laevis (Miiller) 








172 


Agriolimax caruanae Pollonera 








175 


LI MAX Linnaeus 








180 


Limax tenellus Miiller . 










180 


Limax flavus Linnaeus . 










184 


Limax drier eoniger Wolf 










187 


Limax maximus Linnaeus 










191 


LEHMANNIA Heynemann 








193 


Lehmannia marginata (Miiller) 








194 


Greenhouse Aliens ..... 








197 


Lehmannia poirieri (Mabille) . 














197 


Limax nyctelius Bourguignat . 














200 


Fossil Species .... 














202 


Limax modioliformis Sandberger 














202 


Anatomical Terms 














. 203 


Distribution .... 














205 


References .... 














217 


Index ...... 














224 



INTRODUCTION 

No comprehensive work on British slugs has appeared since John William Taylor's 
classic monograph (1902-07). Besides full information on their nomenclature, 
systematics and biology, this contains excellent coloured plates, depicting numerous 
varieties of the eighteen species of slugs which he recognized as British. Besides 
much information on these species which has become available since 1907, five 
more species, a subspecies, and two aliens established in British greenhouses must 
be added to the list. Taylor omitted Arion lusitanicus (p. 135), although it had 
already been recorded from Britain, and it has since reappeared. Milax budapestensis 
(p. 160), another species of Milax provisionally referred to M. insularis (p. 156) 
and Agriolimax caruanae (p. 175) have been recorded ; Agriolimax reticulatus (p. 164) 
has been shown to be specifically distinct from A. agrestis, and Arion ater rufus 
(p. 145) to be subspecifically distinct from A. ater. The alien species Lehmannia 
poirieri (p. 197) and Limax nyctelius (p. 202) have been found in Britain, but only 
in greenhouses. Two names used by Taylor must be changed : Arion circumscriptum 
to A. fasciatus (p. 127) and Limax arborum to Lehmannia marginata (p. 194). 

Two more recent works also deal with all British slugs, though less completely. 
Ellis (1926) gives monochrome figures but only occasional references to internal 
anatomy ; while Quick (1949) also gives monochrome figures, and information 
relating mainly to identification in the field. Among more general works, KunkeJ 
(1916) on the biology of pulmonates, 0kland (1923) on the Arionidae of Norway, 
and Boycott (1934) on the habits of British land molluscs may be especially mentioned. 
Much information on courtship and copulation is given by Gerhardt (1933, 1935, 
1939 and 1940), on the seasonal abundance and growth of slugs in gardens and fields 



BRITISH SLUGS 107 

by Barnes & Weill (1945, with good coloured plates of garden species), and on their 
ecology by Thomas (1944) . For full synonymies, reference should be made to Kennard 
& Woodward (1926), and also to Taylor (1902-07) and Hesse (1926). Only the more 
important synonyms are quoted below. 

The group of naked pulmonates termed slugs is highly polyphyletic in origin. 
British slugs belong to the three families Testacellidae, Arionidae and Limacidae, 
which have evidently been derived from shelled Oleacinids, Endodontids and Zonitids 
respectively. 

A rrangement 

Information is given in the same sequence for each species (except that the shell 
is described before the external variations for Testacella) under the following headings, 
any of which may be omitted. 

Notes Radula 

First British record Alimentary system 

Diagnostic features Reproductive system 

External appearance Spermatophore 

External variation Mating 

Shell Development 

Pallial organs Behaviour 

Pedal gland Ecology 

Retractor muscles Distribution 

Nervous system Fossil record 

Jaw Material examined 

The anatomical terms used in the descriptions are explained by means of a glossary 
and semi-diagrammatic figures on p. 203. Distributions in the British Isles are shown 
on vice-comital maps (p. 205), as well as being briefly described under " distribution ". 

A cknowledgments 

I wish to thank the following. The Trustees of the British Museum for financial 
assistance, and Dr. H. W. Parker, C.B.E. for access to the zoological collections 
of the British Museum (Natural History). Mr. A. E. Ellis for preparing the maps. 
M. J. M. Gaillard for information and specimens from the Museum National d'His- 
toire Naturelle, Paris. Mr. I. C. J. Galbraith for editorial assistance. Dr. H. Loh- 
mander for information on Scandinavian slugs. Dr. O. E. Paget for information 
from the Naturhistorisches Museum, Vienna. Dr. W. J. Rees for valuable advice. 
Mr. H. O. Ricketts for much help in connection with material in the B.M. (N.H.), 
and for his care in preparing the bibliography and index. The late Mr. Hugh Watson 
for much helpful correspondence. The many correspondents who have helped from 
time to time by providing specimens or information, and especially Dr. W. Adam, 
Dr. C. O. van Regteren Altena, Mr. D. Aten, Dr. H. F. Barnes, Dr. A. J. Cain, Mr. 
A. E. Ellis, Dr. L. Forcart, Dr. W. O. Gregg, Dr. P. Makings, Dr. A. D. J. Meeuse, 
Dr. Nils Odhner, Dr. A. W. Stelfox, Mr. Abu Teira, Mrs. M. Vincent, and Dr. M. H. 
Williamson. 



io8 BRITISH SLUGS 

NATIVE AND NATURALIZED SPECIES 

Order STYLOMMATOPHORA 
Pulmonate mollusca bearing eyes at the apex of retractile tentacles. 

Superfamily Oleacinacea 

Jaw rudimentary or absent, radular teeth aculeate, shell turreted or auriform. 
Carnivorous. 

Family Testacellidae 

Shell small, auriform, situated at the posterior end of the body, covering the pallial 
organs. The upper tentacles (ommatophores) are not bulbous at the apex. There is 
no caudal mucus gland. A pair of lateral grooves with side branches directed 
obliquely upwards and forwards, and downwards and forwards, arises from a peri- 
pallial groove. The pedal mucus gland lies free in the body cavity. The cerebral 
commissure is very short, and the cerebro-pleural and cerebro-pedal commissures 
long. There is no jaw. The buccal bulb is enormous. The long narrow radular 
teeth, barbed at the apex, are disposed in transverse V-shaped rows, and a central 
tooth may be absent. The stomach is feebly developed and the anus lies just within 
the pulmonary cavity posteriorly. The kidney is without a secondary ureter. An 
osphradium is said to be present in a groove in the floor of the mantle cavity (Plate, 
1891). The vagina is without accessory glands, and the right ocular retractor muscle 
passes forwards between the penis and vagina. Testacella is carnivorous, feeding on 
earth-worms and occasionally on other slugs, and is largely subterranean in habit. 

Genus TESTACELLA Draparnaud, 1801 

Type species Testacella haliotidea Draparnaud, 1801. 

The sole genus, with the characters of the family. Through adaptation to a life 
spent largely underground, and a diet of earth-worms, the structure of Testacella 
has been profoundly modified. The relatively large shell and mantle cavity of other 
snails have been greatly reduced in size, and the body anterior to these structures 
elongated to accommodate the enormous buccal bulb with its powerful muscles 
and barbed radular teeth. The backwards shift of the pallial region is accompanied 
by a clockwise rotation of 180 , so that the heart and pericardium lie on the right 
side of the kidney with the ventricle in front of the auricle. The normal backwardly- 
directed loop of the intestine is obliterated, and the rectum runs backwards instead 
of forwards to the anus (beside the pulmonary orifice, at the rear). 

It is convenient to gather together the available information on the life history 
of all the three British species of Testacella. A recent paper by Stokes (1958) gives 
further information, supplementing that of Barnes & Stokes (1951). 

Mating 

Little is known, but Gerhart (1940 : 579) reports that while unpacking a container 
he found a pair of T. sctttulum in coitus in the moss, and accidentally separated them. 



BRITISH SLUGS Tog 

The penes were bluish-white, translucent, elongated, dorso-ventrally flattened 
cones, withdrawn from the adjacent vaginal openings. No appendix was visible, 
and the penes were of the simplest type occurring in land pulmonates. They quickly 
disappeared within the owners' bodies, and no more was to be seen. Barnes & 
Stokes (195 1) found that the coitus of T. haliotidea and T. scutulum took place 
under the surface of the soil in their containers, the animals lying head to head : 
they do not mention any visible everted organs. Chemin (1939a and b) reports 
self-fertilization in T. haliotidea isolated when young. 

Development 

The eggs are ellipsoidal, with a white calcareous shell which becomes yellow, 
and are laid underground. Those of the three British species are distinguished 
only by differences in size -.5x4 mm for T. maugei, 7x4 mm for T. haliotidea, 
and 4x3 mm for T. scutulum. Barnes & Stokes confirm the old statement that 
they explode when placed on a warm hand. The eggs of T. maugei are laid in groups 
of 8-15 during the summer months, those of T. haliotidea and T. scutulum in 
captivity (Barnes & Stokes, 195 1) in batches of 2-14 in March, 54-95 days after 
mating. Taylor (1902-07) records the time of development as 30-35 days for 
T . maugei, 20-36 days for T. scutulum and (quoting Gassies, 1849) 10-22 days for 
T. haliotidea. However, Barnes & Stokes found that embryonic development takes 
much longer than this for the latter two species : 144-201 days, or five to six months. 

Like all our slugs and some, perhaps all, of the shell-bearing Stylommatophora 
except Succinea (which lacks a caudal vesicle), Testacella exhibits in the middle and 
late stages of development a haemocoelic cephalic vesicle protruding beneath the 
mantle over the head, containing part of the viscera ; and a caudal haemocoelic 
vesicle projecting beyond the tail, which contains only haemocoelic fluid and cor- 
puscles. The caudal vesicle varies in shape from genus to genus (see Text-fig. 18). 
In Testacella it is a flattened bilobed or broadly cordate sac (Text-figs. 1 d and 18 c). 
The two vesicles pulsate, expanding and contracting alternately so that haemocoelic 
fluid is driven back and forth, at a rate depending on the temperature. The vesicles 
may have an excretory or a respiratory function. The cephalic vesicle is withdrawn 
with its contents into the body, and shortly before hatching the caudal vesicle 
also shrinks and disappears. In Testacella, with its posterior mantle, the cephalic 
vesicle does not extend over the head (at any rate in the later embryonic stages). 

Behaviour 

It has been noted (Adams, 1941) that dust sprinkled on Testacella leads to copious 
discharge of fluid which washes it away. Another phenomenon does not seem to 
have been recorded. When lightly irritated, Testacella maugei partially contracts, 
and sometimes a forcible discharge of air and froth occurs in the middle line from 
under the front of the mantle edge, with a distinct high-pitched hissing sound. 
Presumably a communication to the lung cavity exists, and air is expelled by closure 
of the pulmonary orifice and muscular contraction. No duct is apparent on gross 
dissection, but serial section of the area show spongy tissue leading to the anterior 



no BRITISH SLUGS 

expansions of the pulmonary cavity, through which air might be expelled. In its 
underground travels, or if dragged into a burrow by an attacked earth-worm, it is 
easy to imagine that earth could become impacted under the mantle border, and 
that the forcible expulsion of air and fluid would serve to dislodge it. 

Ecology 

Mostly to be found in gardens, allotments and other cultivated ground. During 
the day Testacella remains underground, retreating deeper in dry weather but some- 
times hiding under stones and herbage in damp weather ; while at night it crawls 
freely on the surface. In the winter it hibernates in a strongly contracted state, 
in an underground cell. On slowly approaching a worm, the slug shoots out its 
radula and impales its prey at one end or in the middle, and engulfs it within a few 
minutes. Another worm may be eaten after a day or two. The faeces are very small 
in bulk and contain discarded radular teeth, fragments of worm cuticle and setae, 
and sometimes small white bodies which may be from the calcareous oesophageal 
glands of worms. Testacella is also said to eat other slugs. 

Testacella maugei Ferussac, 1819 

Testacella maugei Ferussac, 1819 : 94, pi. 8, figs. 10 and 12. Teneriffe. 
Testacella aurigaster Watson, 191 5 : 221 (from Layard MS.). 

Notes 

The type was found in 1796 by M. Mauge ; who died before publishing an account. 
According to Turton (1831 : 27), it was sent in 1812 from Dr. Leach at Bristol to 
Ferussac. T. aurigaster refers to this species, introduced into South Africa (Watson, 
1915 : 221). 

First British record 
By Ferussac (1819). 

Diagnostic features 

Pear-shaped appearance (body widest behind) ; shell large (14 X 7 mm) ; two 
median lines of dorsal tubercles conspicuous, especially in the young ; lateral 
grooves wide (5 mm) apart at origin. Buccal bulb lacks lateral retractors ; cephalic 
retractors arise asymmetrically, from left side. Penis swollen posteriorly, lacks 
the flagellum of T. haliotidea ; penial retractor arises from body-wall about two- 
thirds of the way back ; vas deferens convoluted near free oviduct ; spermatheca 
duct long and slender, swollen proximally. 

External appearance (PL 1, fig. 13) 

Length when extended, 6-10 cm. The most bulky part of the body lies behind 
the centre. The lateral grooves are separated by about 5 mm at their origin from 
the peri-pallial groove, and each gives rise to about fourteen dorsal and ventral 



BRITISH SLUGS in 

grooves directed obliquely forwards, which subdivide to produce the reticulate 
surface pattern. The row of tubercles on each side of the median dorsal line is 
more prominent than the other dorsal rows, especially in the young. The upper 
tentacles, bearing the small black eyes, are not bulbous at the apex, and the lower 
tentacles are short. The true pedal groove is only distinct for a few millimetres at 
the front of the foot fringe, but a peripodial groove above this runs to the tail. 
There is no caudal mucus gland. When crawling on a glass plate, the yellow sole 
is seen to be obscurely tripartite. The mucus is clear and viscid, drawing out into 
thin threads. The body is usually some shade of brown more or less sprinkled with 
darker spots, paler on the sides, and the sole is yellow or pink. The mantle is speckled 
with black, and the paler margin slightly overlaps the shell. The posterior respiratory 
orifice is bounded by the right and left mantle lobes. 

Shell (Text-fig. i a) 

The shell measures about 14 X 7 mm, and has a brown periostracum, often 
abraded. It is more convex than the shells of the other two species, and the columella 
does not show the same callous enlargement at the hinder end. The crescentic 
muscle scar is visible on the under surface. 

External variation 

Variety aperta Taylor (1902) has the shell wider and flatter than normal. Varieties 
albina, griseonigrescens, viridans and griseorubescens Gassies & Fischer (1856), 
nigra Collinge (1898) and aurea Taylor (1902) are colour varieties, whose charac- 
teristics are suggested by their names. 

Pallial organs (Text-figs. 2 d and f) 

The respiratory cavity, bounded by the right and left mantle lobes, leads to the 
pulmonary chamber, which is bounded by spongy vascular tissue. Anterior to 
this, the chamber is prolonged forwards as thin-walled air sacs projecting into the 
visceral cavity. The somewhat quadrate kidney, opening without a secondary 
ureter at the back of the chamber, lies above and in front of the pulmonary chamber. 
The pericardium lies to the right of the kidney and the ventricle is anterior to the 
auricle, and both branches of the aorta run forwards. The rectum runs obliquely 
in the pulmonary chamber to terminate in the anus just above and on the left of 
the respiratory orifice. The kidney projects over the apex of the left liver lobe. 
An osphradium is represented by a ridge of cells in the floor of the respiratory chamber, 
and is innervated from the right visceral ganglion (Plate, 1891). 

Retractor muscles (Text-fig. 1 e) 

Both the cephalic retractor muscles arise from the body-wall on the left side. 
The right ocular retractor passes between the penis and the vagina, and a branch 
takes the same course to the side of the head. The slender penial retractor arises 
from the right body wall about two-thirds of the way back, and not from the posterior 
end as it does in T. haliotidea and T. scutulum. 



BRITISH SLUGS 




Fig. i. Testacella. Reproductive organs (x 3-3), shell (x 2). A. T. maugei — shell 
below. B. T. haliotidea — shell right, below. G. T. scutulum — shell right, 

above. 

D. T. scutulum — advanced embryo ( x 6-7) showing cephalic vesicle in front of 
shell and caudal vesicle posteriorly. E, T. maugei — buccal bulb and ocular 

retractors. 



BRITISH SLUGS 



H3 



Nervous system (Text-fig. 2 e) 

The cerebral commissure is very short, and the cerebro-pleural and cerebro-pedal 
commissures are long to embrace the large buccal bulb. The buccal ganglia are 
large. The five visceral ganglia are close together, but not fused. 

Radula (Text-fig. 2 a) 

In each row, sixteen aculeate barbed teeth on each side of a small median tooth 
increase in size up to the penultimate one, while the sixteenth is much smaller. 
There are about fifty V-shaped rows. 




B 








Fig. 2. Testacella. Representative radular teeth ( x 67). A. T. mmtgei — showing 

central tooth. B. T. haliotidea — showing absence of central tooth. C. T. scu- 

tulum — showing absence of central tooth. 

T. maugei. D. Posterior end of body ( X 3-3) with shell removed and right 

mantle lobe reflected to expose respiratory orifice. E. Central nervous system, 

showing buccal ganglia (black) posterior to cerebral ganglia when animal is contracted. 
F. Pallial region, viewed as if shell were transparent. 



ii 4 BRITISH SLUGS 

Alimentary system 

The buccal bulb, which is large but does not extend as far back as in T. haliotidea 
and T. scutulum, lacks lateral muscles but has a bundle of from two to four pairs 
of powerful muscles at the posterior end, attached to the diaphragm towards the 
left of the mid-line (Text-fig. i e). The small stomach is scarcely wider than the 
oesophagus and intestine. It receives the right and left liver ducts, and the left 
liver lobe forms the apex of the visceral mass. The intestine makes one forwardly- 
directed loop, and the long rectum runs backwards to the anus, close to the respiratory 
orifice. 

Reproductive system (Text-fig. i a) 

The reproductive organs are simple. The pale ovotestis lies at the apex of the 
visceral mass between the right and left lobes of the liver. The slightly sinuous 
hermaphrodite duct becomes slender and folded before entering the long narrow 
albumen gland, and bears a small seminal vesicle. The convoluted vas deferens, 
arising from the yellow ribbon-like prostate, runs forwards beside the free oviduct 
and then turns backwards, beneath the right ocular retractor to the apex of the 
penis. The penis is about 9 mm long, slender in front and dilated posteriorly and 
often folded over forwards. The mucous membrane is papillate at the apex and 
longitudinally ridged beyond, and there is no penial papilla. There is no caecum 
or flagellum. The penial retractor, arising from the right body wall about two- 
thirds of the way back, is inserted at the apex of the penis. The atrium is short 
and opens below the right upper tentacle. The free oviduct and vagina are wider 
than the penis. The long slender spermatheca duct is dilated at its origin from the 
oviduct, and enters the spermatheca sac laterally. The mucous membrane of the 
duct is longitudinally ridged, the ridges being more prominent in the bulbous part 
of the duct. 

Mating, Development, Behaviour, Ecology 

See pp. 108-110. The eggs measure 5x4 mm. 

Distribution (Map 1) 

T. maugei is the most westerly of the three British species of Testacella, known 
from Tangier, south-western Spain, Portugal, Brittany, Dieppe and the Atlantic 
isles (and introduced into North America and South Africa) . In England and Wales 
it occurs west of a line from Cheshire to Middlesex, and in Ireland south-east of a 
line from Dublin to Cork. It is probably truly native only in the south-west. 

Fossil record 

Holocene and Recent. 

Material examined 

Living specimens from Singleton Park Botanical Gardens, Swansea, Glamorgan. 
Spirit material from various British localities, the Azores and Madeira. 



BRITISH SLUGS 115 

Testacella haliotidea Draparnaud, 1801 

Testacella haliotidea Draparnaud, 1801 : 99. ? Montpellier. 
Testacella earopea de Roissy, 1805, 5 : 252. 

First British record 

By Lukis (1834), fr° m Guernsey. 

Diagnostic features 

Less bulky posteriorly, when extended, than T. maugei ; shell much smaller, 
(7x5 mm), columella somewhat truncate anteriorly, with conspicuous callous 
enlargement posteriorly ; dorsal tubercles not especially prominent ; lateral 
grooves close together at origin. Buccal bulb with powerful lateral retractors ; 
cephalic retractors arise symmetrically. Radula lacks a central tooth, apophyses 
of marginal teeth nearly central. Penis with flagellum and short apical caecum ; 
penial retractor arises from diaphragm at posterior end of body ; spermatheca 
duct short and bulbous. 

External appearance (PI. 1, fig. 10) 

Length when extended, 8-12 cm. The lateral grooves are close together at their 
origin from the peri-pallial groove. Colour usually dull greyish-yellow, with a pale 
sole and foot fringe. 

Shell (Text-fig. 1 b) 

Measures about 7x5 mm, convexly auriform, triangular ; brown periostracum 
usually much abraded. Columella usually slightly truncate anteriorly, and expanded 
to a rounded callus posteriorly. 

External variation 

In var. trigona Gassies & Fischer (1856) the shell is thicker and the right margin 
expanded, in var. major G. & F. the shell is larger (11-5 x 7 mm). The body is 
yellow in var. flavescens Moquin-Tandon (1855), and whitish in var. albina Moquin- 
Tandon. 

Pallial organs 
As in T. maugei. 

Retractor muscles 

The cephalic retractor muscles arise symmetrically on each side of the body-wall 
about two-thirds of the way back, and not from the left side only as in T. maugei. 
The long, slender penial retractor arises dorsally at the hind end of the body. 

Nervous system 1 
As in T. maugei. 



n6 BRITISH SLUGS 

Radula (Text-fig. 2 b) 

The median tooth is vestigial or absent. On each side, eighteen barbed aculeate 
teeth gradually enlarge up to the penultimate one, while the eighteenth is shorter. 
The lateral apophysis is nearly midway between base and apex (especially towards 
the margin of the radula) , while in T. scutulum it is nearer the basal end. There are 
about fifty V-shaped rows. 



Alimentary system 

The large buccal bulb is longer than in T. maugei and has, in addition to the bundle 
of posterior muscles attached to the diaphragm, a series of ten to fifteen pairs of 
lateral muscles attached to the posterior half of the bulb and arising from the left 
body-wall. The relations of the liver lobes, intestine and rectum are as in T. maugei. 

Reproductive system (Text-fig. 1 b) 

Compared with T. maugei, the central portion of the hermaphrodite duct is 
longer and more folded, the penis is slighly enlarged at the entrance of the vas 
deferens, and opposite this point there is a short blunt caecum. There is also a 
flagellum, which varies in length but may be two-thirds as long as the penis, from 
the apex of which a long slender retractor muscle runs to the diaphragm at the 
posterior end of the body. There is no penial papilla, and the mucous membrane 
is papillate. The spermatheca duct is short, wide and bulbous with the mucous 
membrane strongly folded longitudinally, and the sac is oval. 

Mating, Development, Behaviour, Ecology 

See pp. 108-110. The eggs measure 7x4 mm, and so are considerably larger 
than those of T. scutulum. 



Distribution (Map 2) 

T. haliotidea is the most widely distributed species, ranging from the Atlantic 
isles to the Balkans, and from the North African coast to south Scotland. It probably 
occurs in every county in Britain up to Stirling, and in the south-eastern and southern 
counties of Ireland. 



Fossil record 

Holocene and Recent. 

Material examined 

Living specimens from Henley-on-Thames, Bedford and Cambridge. Spirit 
material from Britain, France and Majorca. 



BRITISH SLUGS 117 

Testacella scutulum Sowerby, 1820 

Testacella scutulum Sowerby, 1820 : pi. 159, figs. 3-6. Kennington Road, Lambeth, Surrey. 

Notes 

Taylor (1902-07) considers T. bisulcata Risso (1826) as a variety of this species. 
Plate (1891) and Hoffman (1925), however, consider it to be quite distinct. Plate's 
figure, copied by Hoffman, shows a flagellum from the anterior end of the penis, 
and a tubular connection from the vagina to the apex of the flagellum. This most 
unusual arrangement is said to be confirmed by serial sections. 

First British record 
By Sowerby (1820). 

Diagnostic features 

Resembles T. haliotidea, but differs in the following. Shell smaller (6 X 3-5 mm) 
and flatter ; animal yellow in colour, speckled with brown. Apophyses of marginal 
radular teeth more basal. Penis lacks caecum and flagellum ; free oviduct shorter 
than vagina ; spermatheca duct longer and narrower (but quite different from that 
of T. maugei). 

External appearance 

Length when extended, 8-12 cm. The lateral grooves are said to be confluent 
for about 1 mm at their origin from the peri-pallial groove, but this does not seem 
to be constant. Colour some shade of yellow, more or less sprinkled with brown ; 
sole and foot fringe brighter yellow. 

Shell (Text-fig. 1 c) 

Smaller than that of T. haliotidea (6 X 3-5 mm), and flatter — sometimes even 
concave. The brown periostracum tends to be retained better than in the other 
species. The columella is slightly truncate anteriorly and expanded posteriorly, 
as in T. haliotidea. Young shells of these two species can easily be confused. 

External variation 

Most of the varieties mentioned by Taylor (1902-07) refer to continental forms, 
which may or may not be conspecific with T. scutulum. However, var. aurea Cockerell 
(1885) presumably refers to this species : it is orange-coloured, with brown mottling 
and a bright orange foot sole. 

Pallial organs 

As in T. maugei and T. haliotidea. 



n8 BRITISH SLUGS 

Retractor muscles 

The cephalic retractors arise symmetrically from the body-wall about half-way 
back. The penial retractor arises far back, from the diaphragm. 

Nervous system 

As in T. maugei and T. haliotidea. 

Radula (Text-fig. 2 c) 

As in T. haliotidea, the median tooth is absent and there are eighteen teeth on 
each side of the mid-line ; but in the outer marginal teeth the apophyses are nearer 
the bases. There are about forty-five V-shaped rows. 

Alimentary system 

The buccal bulb has three or four strong muscles from the diaphragm to its 
posterior end, and the hinder half has five to ten pairs of lateral muscles attached 
to the left body-wall. The relations of the liver lobes, intestine and rectum are as 
in T. maugei and T. haliotidea. 

Reproductive system (Text-fig. i c) 

The ovotestis, albumen gland, prostate and oviduct resemble those of T. haliotidea, 
but the free oviduct is relatively shorter and the vagina longer. The spermatheca 
duct is intermediate in length and breadth between those of T. haliotidea and 
T. maugei. The penis is cylindrical, without dilatations or flagellum, and the retractor 
muscle arises far back from the diaphragm. The penial mucous membrane is papillate, 
and that of the spermatheca thrown into longitudinal folds. 

Mating, Development, Behaviour, Ecology 

See pp. 108-110. The eggs are smaller (4x3 mm) than those of the other species, 
but otherwise similar. 

Distribution (Map 3) 

Taylor (1902-07 : 18) apparently treated T. bisulcata Risso (1826) as a variety 
of T. scutulum, so that his account of the latter 's continental distribution is unreliable. 
In the British Isles, its distribution is almost the same as that of T. haliotidea. 

Fossil record 
Recent. 

Material examined 

Living specimens from Bedford. Spirit material from Britain and Corsica. 



BRITISH SLUGS 119 

Superfamily Endodontacea 

Shell spiral depressed, or reduced to a flat plate or mass of granules enclosed 
by the mantle. A supra-pedal groove is present, and a caudal mucus gland may be 
present or absent. The jaw is either crescentic and ribbed, or plaited. Marginal 
radular teeth generally aculeate. The genitalia are simple. 



Family Arionidae 

Slug-like animals with the mantle covering only the anterior part of the body, 
with the respiratory orifice in a slit on the right margin. Shell an oval plate or mass 
of granules except in Binneya, where it is exposed and spiral. The sole is uniform or 
tripartite. The kidney surrounds the pericardium. An epiphallus is present. The 
jaw is ribbed (odontognath) , the central radular tooth tricuspid, lateral teeth 
bicuspid, marginals with short wide basal plates, with long mesocones and with or 
without ectocones. 

The Arionidae require relatively high humidity, and dry barren areas are barriers 
to their dispersal. The family seems to have been derived from Endodontid stock 
with spiral shells, the cephalic and buccal retractor muscles with a common origin 
from the columella, and a penial papilla. In the primitive North American genus 
Binneya, the visceral hump is capped by a spiral Vitrina-like shell, the tail is solid 
and the retractors have a common origin. Various genera show transitions from this 
snail-like condition to that of the most modified subfamily, the Arioninae of the 
western Palaearctic Region. For a discussion of the probable evolution of the other 
subfamilies and genera, see Pilsbry (1948 : 660-665). 

Subfamily Arioninae 

Slugs with a wide foot fringe ; caudal mucus gland present ; the shell is reduced 
to a flattened plate in Geomalacus, and to discrete granules in Avion. The cephalic 
retractor muscles are widely separated at their origin from the posterior border 
of the diaphragm. The stomach apex lies behind the posterior loop of the intestine. 
The ovotestis lies behind all the intestinal loops. There is no penis or vagina, and 
the oviduct, spermatheca and epiphallus enter the upper atrium separately. 

Of the British genera, Geomalacus with its solid shell, anterior reproductive 
orifice, narrower foot fringe, and extreme western distribution, seems to be the more 
primitive. 

Genus GEOMALACUS Allman, 1843 

Type species Geomalacus maculosus Allman, 1843. 

Genital orifice below and behind the right upper tentacle ; atrium greatly elong- 
ated ; genital retractor muscle arising from the dorsal body-wall at the extreme 
posterior end of the body and inserted into the spermatheca duct ; shell a compact 
oval calcareous mass. 

zool. 6, 3. q 



BRITISH SLUGS 



G. maculosus is the only member of this genus occurring in the British Isles, though 
one or two more are found with it in the Iberian peninsula. 



Geomalacus maculosus Airman, 1843 
Geomalacus maculosus Allman, 1843 : 851. Lough Caragh, Co. Kerry, Ireland. 
Notes 

The types were found by W. Andrews of Dublin in 1842. 

First British record 
By Allman (1843). 

Diagnostic features 

Distinguished from species of Arion, externally by the anterior situation of the 
reproductive orifice, narrower median area of the sole, and maculate colour pattern ; 
and internally by the solid oval shell, greatly elongated atrium, and long genital 
retractor arising far back from the mid-dorsal line of the body-wall. 

External appearance (PL 1, fig. 1) 

Length when extended about 5-5 cm, but occasionally up to 9 cm. The granulose 
mantle occupies about one-third of the body, with the respiratory orifice a little 
in front of the centre of the right border. The reproductive orifice is nearer to the 
right upper tentacle than to the respiratory orifice. The supra-pedal groove is 
deep and distinct and the pedal groove indistinct. The caudal mucus gland is a 
transverse slit. The body shows about twenty-five rows of polygonal tubercles 
(Text-fig. 3 c). The colour is grey, brown or nearly black, irregularly marked with 
yellow or white maculations more or less developed. The young have two longitudinal 
bands on each side, which tend to become obscured in the adult, but sometimes 
remain. In contrast to Arion, Geomalacus sometimes rolls up like the woodlouse 
A rmadillidium . 

External variation 

Var. fasciata Cockerell (1892) retains the banding characteristic of the young — 
animal white or whitish with black or brown markings : mantle with dark marbling 
and lateral bands, body with a dark subdorsal band along each side. Charles 
Oldham bred in captivity a crimson form, var. rutilans Oldham (1 942ft), which has 
not been observed in the wild. The other varieties depend merely on the relative 
development of dark pigment on a pale ground : var. allmani Heynemann (1873), 
dark brown or grey with whitish maculations ; var. verkruzeni Heynemann, grey, 
darker dorsally, with whitish maculations ; var. andrewsi Mabille (1867), whitish, 
overspread with blackish spots. 



BRITISH SLUGS 121 

Shell (Text-fig. 3 1) 

Variable, but usually solid, oval, convex above and concave below, measuring 
about 4x3 mm. 

Pallial organs (Text-fig. 3 b) 

The kidney completely surrounds the pericardium, and the cavity is occupied 
by lamelliform folds. The ventricle lies behind the auricle. The anterior branch of 
the aorta follows the usual course, arching over and passing beneath the anterior 
intestinal loop, and running forwards between the visceral and pedal ganglia to 
supply the anterior end of the body. The posterior aorta runs backwards to the 
digestive and genital systems. 

Retractor muscles (Text-fig. 3 b) 

The right and left cephalic retractor muscles are widely separated at their origin 
from the diaphragm, behind the kidney. The buccal retractor arises from the dorsal 
body- wall, a little further back and slightly to the right of the middle line. The right 
tentacular branch lies to the left of the oviduct and epiphallus, and does not pass 
between them. 

Nervous system 

The cerebral commissure is of medium length. The cerebral ganglia give rise to 
nerves to the head and lips, the peritentacular nerves, the combined ocular and 
olfactory nerve, the nerve to the lower tentacle, and on the right side to the genital 
nerve supplying the epiphallus and vas deferens. The right pleural, visceral and 
abdominal ganglia are fused together, but just distinguishable, and the same is 
true of the left pleural and the left visceral ganglia. The pleuro-pedal commissures 
are very short. The buccal ganglia, as well as supplying the buccal bulb, send a 
nerve to the salivary glands. 

Jaw (Text-fig. 3 d) 

The jaw is about 3 mm across, dark brown, arcuate, and crossed by eight or ten 
broad ribs which slightly denticulate the upper and lower margins. The number 
of ribs may be reduced by fusion, and there is sometimes a slight median projection. 

J Radula (Text-fig. 3 e) 

The radula is about 8 mm long. There are usually about 200 rows of teeth, but 
the number varies considerably. The central tooth has very small ectocones. The 
lateral teeth, about twenty in number, have a large mesocone, a small ectocone, 
and no endocone. The ectocone gets smaller in the outer lateral teeth. In the marginal 
teeth it descends to the base of the mesocone, and gets larger again as the margin 
of the radula is approached. 



BRITISH SLUGS 








ko SI 

99 



ko> 





H 



Fig. 3. Geomalacus maculosus. A. Reproductive organs (x 2-7). B. Head 

and mantle region (x 1*7) with mantle reflected to the right showing kidney, peri- 
cardium, rectum, terminations of genital and alimentary systems, origin of pharyngeal 
and left ocular retractors cut short. G. Polygonal body- tubercles. D. Jaw 

(x 13). E. Representative radular teeth ( x 330). I. Shell. 

Avion. Representative radular teeth (x 330). F. A. ater. G. A. lusi- 

tanicus. H. A. subfuscus. 



BRITISH SLUGS 123 

Alimentary system 

The oesophagus, after passing through the nerve ring, enlarges to a crop, merging 
into the stomach which forms the most posterior part of the alimentary canal. 
The narrower intestine passes far forwards (to the level of the kidney) and then 
turns back, but does not reach the stomach before it runs forwards again, as the 
rectum, to end in the anus in the posterior lip of the respiratory orifice. The short 
ducts of the anterior and posterior lobes of the liver open at the junction of stomach 
and intestine. 

Reproductive system (Text-fig. 3 a) 

The somewhat raspberry-shaped brown ovotestis lies between the upper and lower 
lobes of the liver, at the apex of the stomach. The pale hermaphrodite duct passes 
forwards, becoming wider and folded, towards the linguiform albumen gland. It 
then becomes narrower, runs backwards between the lobes of the albumen gland, 
and turns sharply forwards, forming a " claw" or seminal vesicle. The common 
duct is much folded, the prostatic portion showing as a yellow ribbon. The narrower 
free oviduct enters the lower end of the atrium ; its mucous membrane is thrown 
into fine longitudinal folds. The lower atrium shows internally a complicated 
mass of large sinuous folds, and is also remarkable in having a long slender diverticulum 
with circular folds, extending backwards as far as the albumen gland. The short 
duct of the globular spermatheca enters the apex of this diverticulum. The vas 
deferens runs backwards from the free oviduct towards the apex of the atrial diverti- 
culum, to enter the wider epiphallus which is very long and much coiled, the coils 
being bound together by connective tissue. The epiphallus enters the atrial diverti- 
culum close to the spermatheca duct. A long slender retractor muscle, arising from 
the mid-dorsal line at the posterior end of the body, is inserted at the neck of the 
spermatheca and apex of the diverticulum. The mucous membrane of the epiphallus 
shows longitudinal rows of papillae, and that of the spermatheca longitudinal 
folds. A membrane with a series of muscular bands extends from the atrial diverti- 
culum to the diaphragm, and similar bands from the lower atrium to the body- wall. 

Mating 

This has not been recorded. 

Development 

The ellipsoidal eggs are said (Taylor, 1902-07 : 256) to be relatively large, from 
8-5 x 4-25 to 6 x 4 mm, and to be white when laid, turning brown. They are laid, 
in clusters of from 18 to 24, in July and August, and hatch in from six to eight 
weeks. The young, which mature in the following summer, show two dark bands on 
each, side which fade during growth. 

Behaviour 

When irritated, it can roll up like the woodlouse Armadillidium (Taylor, 1902-07 : 
257)- 



I2 4 BRITISH SLUGS 

Ecology 

G. maculosus lives on the bare mountain slopes of Old Red Sandstone in Cork and 
Kerry, where it browses on the algae, mosses and liverworts which coat the boulders. 
In captivity it readily feeds on lettuce and carrot, and has been observed to attack 
small snails such as Vitrina pellucida. 

Distribution (Map 4) 

Portugal, north-western Spain, and Vennes (France). In Britain it is confined 
to south-western Ireland, in the counties of Cork and Kerry. 

Fossil record 
Recent. 



Material examined 

Preserved specimens from Glengariff, Co. Cork (August 1931), and other localities 
in Cork and Kerry. 



Genus ARION Ferussac, 1819 

Type species Arion empiricorum Ferussac (— Limax ater Linnaeus, 1758). 

The diagnostic features of the genus are those of the subfamily Arioninae. 
Differences from Geomalacus are : reproductive orifice immediately below posterior 
lip of respiratory orifice ; upper atrium not prolonged backwards as a diverticulum ; 
genital retractor short, arising from diaphragm close to left cephalic retractor, and 
inserted into both spermatheca duct and free oviduct ; shell reduced to discrete 
granules ; median area of sole wider. 

A . intermedins and A . fasciatus seem to be less advanced than the other species, 
in the absence of a ligula, the limited eversion of the gentalia during coitus (in 
A. fasciatus at least), and the slightly more anterior position of the reproductive 
orifice in A. fasciatus. A. hortensis differs strikingly from A. fasciatus and from the 
larger species, but agrees with A. subfuscus and A. lusitanicus, in having the ligula 
(or its functional equivalent) in the proximal part of the oviduct. A. lusitanicus 
is in some respects intermediate between A . subfuscus and A . ater ; while the latter, 
in which the banding pattern disappears completely in the adults, is probably the 
least primitive. 

The various subgenera and sections that have been proposed are not satisfactorily 
delimited. Hesse (1926 : 64-69) uses the following subgenera : Lochea Moquin- 
Tandon (1855), for A. ater and A. lusitanicus (though the genitalia of the latter are 
much more like those of A . subfuscus) ; Mesarion Hesse, for A . subfuscus ; Carinarion 
Hesse, for A. fasciatus ; Kobeltia Siebert (1873), for A. hortensis ; and Microarion 
Hesse, for A . intermedins. 



BRITISH SLUGS 125 

Arion intermedins Normand, 1852 

Avion intermedins Normand, 1852 : 6. Valenciennes. 
Arion minimus Simroth, 1885 : 237, pi. 7, fig. 41. 

First British record 

By Alder (1848 : 124), as Arion flavus. 

Diagnostic features 

Small size, tubercles assume a conical shape with translucent apices when animal 
contracts, yellow mucus accumulates at each end of foot ; reproductive orifice 
slightly anterior to respiratory orifice. 

External appearance (PI. 1, fig. 7) 

This is the smallest British Arionid, about 2 cm long when crawling. Its colour 
varies from yellow to some shade of grey, with the head darker grey or black, and 
there may or may not be body and mantle bands. All these colour varieties may be 
found living together. A. intermedins is easily distinguished from the young of 
larger species by the form of its tubercles, which stand up in little conical eminences 
with translucent tips when the animal contracts — hence the popular name Hedgehog 
Slug. The pair of mantle bands together form a lyre-shaped figure, and the right 
band arches over the respiratory orifice and sends a branch down below and behind 
it. The sides of the body are paler than the back, the foot fringe is yellowish-grey, 
and just above it there is a row of black dots towards the anterior end. The sole is 
yellowish-grey, and yellow mucus tends to accumulate at each end of it. The genital 
orifice is immediately below the slit of the respiratory orifice, and thus is further 
back than in A . fasciatus but slightly further forward than in the other species. 

External variation 

Var. normalis Moquin-Tandon (1855), yellow with blackish head and tentacles ; 
var. plumbea Collinge (1892&), dark grey ; var. brunnea Taylor (1902-07), maroon- 
brown with darker bands ; var. alba Taylor, almost white. 

Shell 

The shell granules are not specifically distinguishable. In calcareous sites they 
may be aggregated into an irregular mass. 

Pallial organs 

As in A. ater (p. 139). 

Retractor muscles 

As in A. ater, except that the cephalic retractors do not arise at the same level, 
but with the right a little further forward than the left ; while the genital retractor 



126 BRITISH SLUGS 

sends a branch to the junction of oviduct and atrium, as well as to the neck of the 
spermatheca. 

Nervous system 
As in A . ater . 

Jaw 

About i mm across, arcuate and crossed by seven to ten widely-spaced ribs. 
Of the usual Arionid shape, and so not like Taylor's figure (1902-07, fig. 248). 

Radula 

About 2 mm long, with about 120 transverse rows of teeth. The base of the 
mesocone of the central tooth is more constricted than in the other species. There 
are about sixteen lateral and sixteen marginal teeth, each with mesocone and 
ectocone : they do not have endocones. Formula C.16.16 x 120. 

Alimentary system 

As in A. ater, except that the visceral mass is less twisted. 

Reproductive system (Text-fig. 4 h) 

The deeply pigmented ovotestis lies between the lobes of the liver, and is partly 
exposed on the surface of the visceral mass. The central part of the pale hermaphro- 
dite duct is wider and somewhat sinuous, then narrows and runs back on the short 
linguiform albumen gland and bends sharply forwards, making a " claw " or seminal 
vesicle. The sperm-oviduct is short, and the free oviduct enters the very short 
upper atrium without a ligula. The mucous membrane is thrown into five or six 
low longitudinal folds. The lower atrium is about 2 mm long and smooth internally. 
The spherical spermatheca is connected to the upper atrium by a very short duct, 
much dilated at its base. The vas deferens is about 2 mm long, and the epiphallus 
about the same length. 

Spermatophore 
Unknown. 

Mating 

Not recorded. 

Development 

The ellipsoidal eggs are relatively large (2 X 1-5 to 2 X 2 mm), opaque, white 
and dotted with calcareous spots. They are laid in the summer and autumn, and 
hatch in about three weeks. The young at birth are 4 mm long, and easily recogniz- 
able as miniatures of the adults. 



BRITISH SLUGS 127 

Ecology 

Common in the moist ground-litter of woods and copses, beneath logs, and at the 
roots of grass in open fields. Often found feeding on toadstools in the wild, and in 
captivity readily eats carrot, lettuce and oatmeal. 

Distribution (Map 5) 

Portugal, Spain, France, northern Italy, Switzerland, Germany, Scandinavia, 
Russia, Azores (and introduced into New Zealand, Polynesia and North America). It 
probably occurs in every vice-county in the British Isles, including islands north- 
wards to the Orkneys and Shetland. 

Fossil record 

Although shell granules of Avion are not recognizable with certainty, they are 
recorded from the Cromerian onwards. They cannot be assigned specifically. 

Material examined 

Living specimens from many parts of England and Wales. Spirit material from 
Britain, France, Norway and Tahiti. 



Arion fasciatus (Nilsson, 1822) 

Limax fasciatus Nilsson, 1822 : 4. ? Sweden. 
Avion circumscriptus Johnston, 1828 : 76. 

Notes 

Collinge (1892 b) reinstated Nilsson's name, though it has not been adopted by 
other British authors. Lohmander (1937) also concluded that fasciatus is the valid 
name, though he continued to use circumscriptus. Nilsson's description of his species 
leaves no doubt that it is the A. circumscriptus of most authors, and Collinge's use 
of fasciatus prevents any action being taken to conserve circumscriptus. 

First British record 
By Johnston (1828). 

Diagnostic features 

Body bell-shaped in transverse section when contracted, mid-dorsal line of 
tubercles pale and raised, reproductive orifice in front of respiratory orifice, right 
mantle band arches over respiratory orifice, sole opaque porcelain white ; sperma- 
theca with short stout duct and long tapered sac, spermatophore coiled anteriorly. 

External appearance (PL 1, fig. 5) 

Length when crawling about 3-3*5 cm. Easily distinguished from A. hortensis 
by the opaque white sole, and by the bell-shaped transverse section of the body 



128 BRITISH SLUGS 

when contracted (caused by splaying-out of the lower part). The reproductive 
orifice is further forward than in our other species, being a little in front of the slit 
of the respiratory orifice. Body and mantle some shade of grey, becoming paler 
towards the foot and with a dark lateral band on each side. The mantle bands form 
a lyre-shaped figure, and the right band arches over the respiratory orifice without 
sending a branch below it. The bands are set at a higher level than in A. hortensis. 
The tubercles of the mid-dorsal line are larger and paler than those on each side, 
forming a keel (hence the sectional name Carinarion Hesse, applied to this species) : 
this is often more distinct in the young. The foot fringe is somewhat lineolated, 
especially towards the tail. 

External variation 

Var. grisea Collinge (1892c), light silver-grey with dark bands ; var. flavescens 
Collinge (1892a) (= var. circumscripta 0kland, 1923), light brownish-yellow, with 
dark bands and a yellow zone over the foot. Lohmander (1937) discusses whether 
these and other named forms are varieties or subspecies. 

Pallial organs 
As in A . ater. 

Retractor muscles 

The genital retractor is mainly inserted into the duct of the spermatheca, but 
usually sends a small branch into the middle of the free oviduct. Otherwise as in 
A. ater. 

Nervous system 
As in A . ater. 

Jaw 

Of the usual arcuate shape, crossed by ten to twelve ribs ; about 1-25 mm across, 
coloured yellow to pale brown. 

Radula 

Closely resembles that of A. hortensis, but the marginals often have the small 
ectocone better developed. Length 3-4 mm, formula C.11.23 X 120. 

Alimentary system 

As in A . ater, except that the visceral mass is less twisted. 

Reproductive system (Text-fig. 4 1) 

The ovotestis is darkly pigmented, the hermaphrodite duct has the usual " claw " 
or seminal vesicle at its entry into the albumen gland, and the sperm-oviduct is 
pinkish in colour, with a broad yellow prostatic ribbon. The free oviduct is rather 



BRITISH SLUGS 129 

long and slender, agreeing with that of A . intermedins in not being dilated terminally 
to contain a ligula as in A. hortensis. The spermatheca has a short stout duct and 
an elongated sac, wide at the base and tapering apically. It varies a good deal, 
sometimes assuming bizarre shapes. The vas deferens is about the same length 
as the epiphallus, which terminates with a slightly marked annular swelling. The 
upper atrium is very small, and contains no ligula. The lower atrium is long, dorso- 
ventrally flattened, and yellow. 

Spermatophore (Text-fig. 4 g) 

A chitinous tube about 4 mm long, with five low ridges of which the most prominent 
is finely toothed. Within the spermatheca, the spermatophore is bent into a U, 
and is swollen for a short distance behind the slender coiled anterior end. On this 
swollen part, Lohmander (1937) describes, but does not figure, a small projection 
which anchors the spermatophore to the mucous membrane of the spermatheca. 
0kland (1923) figures an incomplete spermatophore, and Lohmander (1937) complete 
ones. 

Mating 

Coitus lasts about twenty minutes. In contrast to A. hortensis, A. subfuscus 
and A. ater, there is little or no eversion of the genitalia. 

Development 

The ellipsoidal eggs, 3x2 mm, are yellow or sometimes dark amber, more 
flexible and translucent than those of other species of Arion. They are laid in clutches 
of up to thirty, from May to the autumn, and hatch in four or five weeks (or longer 
in cold weather) . The newly-hatched young are 5 mm long, pale yellowish-grey ; 
not darker dorsally, but with a pale central line of larger tubercles forming a keel 
towards the tail. The lyre-shaped figure formed by the mantle bands is faint in 
front and darker posteriorly. 

Ecology 

A. fasciatus is common amongst dead leaves and moss and under logs and stones, 
in thickets, fields and waste places, and sometimes in gardens. 

Distribution (Map 6) 

Probably every vice-county of the British Isles, and in Iceland (? introduced), 
the Faroes, Scandinavia, France, northern Italy, Rumania, western Russia, ? Spain. 
Introduced into North America. 

Material examined 

Living from many places in Britain. Spirit material from Britain, France and 
Italy. 



130 BRITISH SLUGS 

Arion hortensis Ferussac, 1819 

Avion hortensis Ferussac, 1819: 65, pi. 2, figs. 4-6. Presumably France. 
Avion elongatus Collinge, 1894 : 66, pi. 5A, figs. 1-4. 
Avion coevuleus Collinge, 18976 : 444. 

First British record 
By Gray (1821 : 239). 

Diagnostic features 

Differs from A . fasciatus in the following : body semicircular in transverse section 
when contracted, no enlarged mid-dorsal tubercles, reproductive orifice further 
back, mantle band surrounds respiratory orifice, sole yellow or orange ; spermatheca 
globular, free oviduct longer and proximally more slender, spermatophore hooked 
posteriorly and either discoidal or pointed anteriorly, brown patch on buccal bulb. 

External appearance (PI. 1, fig. 4) 

Length extended 2-5-3 cm - When contracted, the body is semicircular, not bell- 
shaped, in transverse section. The reproductive orifice is below the posterior lip 
of the respiratory orifice. The dark brown or black back is sprinkled with yellowish- 
brown dots, giving a pepper-and-salt appearance, the sole is pale yellow to bright 
orange, and the mucus is yellow. In alcohol, the back becomes bluish-grey and the 
sole loses its orange colour. The lateral bands are placed lower on the body than in 
A. fasciatus, and run along the edges of the mantle, surrounding the respiratory 
orifice instead of arching over it. 

External variation 

Var. grisea Moquin-Tandon (1855), P a ^ e g re Y an< ^ bandless ; var. nigra Moquin- 
Tandon, bands nearly obliterated by general dark pigmentation ; var. rufescens 
Moquin-Tandon, reddish with black bands ; var. caerulea Collinge (1892a), bluish 
with dark bands, and yellow sides. 

Pallial organs 
As in A . ater. 

Retractor muscles 
As in A . fasciatus. 

Nervous system 
As in A . ater. 

Jaw (Text-fig. 7 g) 

About 1 mm across, doubtfully distinct from that of A . fasciatus. 



BRITISH SLUGS 131 

Radula 

Similar to that of A. fasciatus, but the ectocone of the marginal teeth is less 
frequently present. Formula C.i 1.24 X 100. 

Alimentary system (Text-fig. 4 a) 

Visceral mass less twisted than in A. ater. The buccal bulb has a brown shield- 
shaped mark on the upper surface — an apparently good specific character not 
recorded in the literature. 

Reproductive system (Text-figs. 4 b and c) 

The very darkly pigmented ovotestis lies over the apex of the stomach, near the 
posterior end of the visceral mass, and is largely exposed on the surface. The middle 
dilated portion of the hermaphrodite duct is much folded, yellowish- white, and the 
terminal slender part shows the usual " claw " or seminal vesicle where it enters 
the elongated albumen gland. The sperm-oviduct is pale pinkish-brown, with a 
conspicuous yellow ribbon-like prostate. The free oviduct is unusually long and 
slender proximally, and dilated distally before entering the small white upper 
atrium. The vas deferens is short, and the epiphallus about the same length. The 
spermatheca duct is short and stout, with a bulbous base, and the sac is spherical. 
The lower atrium is yellow, with a soft glandular covering. 

As in A. fasciatus and A. intermedins, there is no ligula in the upper atrium. 
However, within the dilated distal end of the oviduct (Text-fig. 4 c), two conspicuous 
longitudinal folds correspond functionally with this structure during coitus (Quick, 
1946). Below the termination of the spermatheca duct is a region of thickened 
mucous membrane in the atrium. In the slugs observed in Swansea (see below), 
the entering spermatophore adhered to this region. The epiphallus terminates in 
a short papillate papilla, which is very conspicuous when the atria are everted in 
coitus (Text-fig. 4E). 

Both British and Dutch slugs show some variation in the folds of the oviduct 
representing the ligula, and in the relative length and thickness of the firm carti- 
laginous part ; but these differences could be due to differences in the degree of 
maturity of the animals, and to the extent of contraction during fixation in alcohol. 

Spermatophore (Text-figs. 4 f and m) 

Two types of spermatophore have been found in slugs from different localities. 
Since the spermatophores of Avion are seldom seen, it seems better to regard the 
slugs bearing the two types as varieties or local races, rather than as distinct species, 
until further anatomical or ecological differences are demonstrated. 

Slugs from Reading and from Zaandam in Holland (collected by Mr. D. Aten) 
have spermatophores in the form of simple chitinous tubes, 6-5 mm long, recurved 
at the slender posterior end and slightly shouldered near the anterior end, smooth 
and without denticulation (Text-fig. 4 F). 

Slugs from a garden in Eaton Crescent, Swansea (Glamorgan), which otherwise 
agree with those from Reading, have the very different spermatophores described 



132 BRITISH SLUGS 

by Quick (1946). These are 5 mm long, shaped like cloves, with the anterior end 
forming a flattened disc with a crenulated edge, and with a ridge carrying about 
fifty-six forwardly pointing denticulations running the whole length of the convex 
side (Text-fig. 4 m). 

Mating (Text-fig. 4 d) 

Mating occurs in April and December, and probably in other months. After the 
usual " following" and licking of mucus, slugs from Swansea (spermatophores as 
in Text-fig. 4 m) curve in a semicircle head to head, and the atria are everted as a 
translucent mass with the lower atria stretched to a narrow yellow ring and the 
lower atria in contact. A long slender mobile club-shaped process is everted, which 
plays over the partner's body in the same manner as the sarcobelum of Agriolimax. 
However, subsequent dissection shows it to be the everted slender proximal part 
of the oviduct, functioning as a stimulatory organ, a modification which appears 
to be unique (Text-fig. 4 e). 

Coitus in Swansea slugs lasts for about an hour and a half. Gerhardt (1935) 
describes the mating of A. hortensis as lasting only twenty-five minutes, and the 
everted masses as small and featureless, in slugs whose spermatophores lacked 
serrations. This suggests that more than one species may be involved, though 
Mr. D. At en (in litt.) states that the Zaandam slugs, having smooth spermatophores, 
evert their oviducts in coitus. 

Development 

The opaque yellow, sticky eggs measure 2-5 X 2 mm, and are laid in clusters 
of up to thirty. A second and third clutch may follow at three-weekly intervals. 
In the winter, the eggs take six or seven weeks to hatch. The newly hatched young 
are 4 or 5 mm long, and differ from those of A . fasciatus in having a darker dorsum 
and central mantle area. The usual body bands and lyriform mantle bands are 
present, and there is no dorsal line of pale conspicuous tubercles. The tentacles 
are violet-brown. 

Ecology 

This species is very common in gardens, allotments and fields, and burrows in 
the ground more than the other species of Avion. It can become a serious pest, 
by eating carrot, potato, etc. It is also frequent in woodland under logs and debris. 

Distribution (Map 7) 

Every county in the British Isles ; Scandinavia and western Europe to western 
Russia, (North Africa). Introduced into North America. 

Material examined 

Living and in spirit from many places in England and Wales, and from Zaandam, 
HoUand. 



BRITISH SLUGS- 133 

Arion subfuscus (Draparnaud, 1805) 

Limax subfuscus Draparnaud, 1805 : 125, pi. 9, fig. 8. France. 
Arion fuscatus Ferussac, 1819 : 65, pi. 2, fig. 7. 

First British record 

Possibly by Johnston (1838), as Arion subflavus (a nomen nudum). Up to about 
1880, however, A. subfuscus was generally considered as a variety of A. ater. 

Diagnostic features 

Adults larger (6—7 cm) than A.fasciatus and A. hortensis, smaller than A. lusitanicus 
and A . ater ; differ from young of large species in smaller tubercles, narrower foot 
fringe, inability to contract to hemispherical shape. Internally differs from A . hortensis 
in relatively shorter proximal, slender, part of free oviduct, longer spermatheca 
duct. 

External appearance (PL 1, fig. 2) 

Length extended, from 5 to 7 cm. The colour is usually yellowish-brown, darker 
on the dorsum and head and on the body bands. On the mantle the right band 
completely surrounds the respiratory orifice. The foot fringe is yellowish and lineo- 
lated more or less distinctly with black, and the sole is pale yellow, sprinkled with 
whitish specks, and obscurely tripartite. The mucus, especially on the mantle, 
is yellow or orange. When contracted, A . subfuscus neither assumes the hemispherical 
shape nor exhibits the rolling reflex (p. 145) of A. ater. 

External variation 

This as usual depends on the relative development of black, red and yellow pigment, 
and the distinctness of the bands : var. rufofusca Draparnaud (1805), rufous with 
black bands ; var. bicolor Moquin-Tandon (1855), red with darker dorsum and no 
bands, foot fringe orange ; vzr.fuliginea Morelet (1845), dark brown, fringe yellowish; 
var. nigricans Pollonera (1887&), black ; var. cinereofusca Draparnaud (1805), 
ashy grey ; var. succinea Bouillet (1835), yellow or orange ; var. pegorarii Lessona 
& Pollonera (1882), quadrifasciate ; var. alba Esmark (1883), whitish. In most 
cases, the yellow pigment is largely contained in the mucus, and when this is removed 
the animal becomes brown or grey, as do alcohol specimens. 

Pallial organs, retractor muscles, nervous system 
As in A . ater. 

Jaw 

Brown, arcuate, about 1-5 mm across, with from ten to sixteen transverse ribs. 



134 BRITISH SLUGS 

Radula (Text-fig. 3 h) 

About 4-5 X 2 mm. About ten more teeth on each side of each row than in 
A. hortensis. Formula C.15.30 x 140. 

Alimentary system 

Similar to that of the other species of Avion, except that the visceral mass is 
rather more twisted than in A. intermedins and A. fasciatus, and less so than in 
A. ater. 

Reproductive system (Text-figs. 6 d and f) 

From the pale brown ovotestis, the greatly convoluted white hermaphrodite 
duct enters the albumen gland, with the usual "claw" or seminal vesicle. The 
common duct is voluminous and folded, the male or prostatic portion yellow and 
the female part pinkish-grey. The first part of the free oviduct is short and slender, 
and the longer distal part dilated. This part contains two prominent longitudinal 
folds of mucous membrane forming the ligula as in A. hortensis. The spermatheca 
duct is bulbous at its origin, then longer and more slender than in A. hortensis, 
and ends in a larger spherical spermatheca sac. The vas deferens and epiphallus 
are very similar to those of A. hortensis, but the terminal annular thickening is 
more pronounced. The upper atrium is small, the yellow glandular lower atrium 
of medium size with a wrinkled mucous membrane. 

Spermatophore (Text-fig. 6 g) 

Very different from that of A . hortensis : about 20 mm lpng, slender anteriorly, 
then dilating, to taper again to an almost filamentous tail. The convex edge is 
finely denticulate. 

Mating 

Mating occurs in March, April and May, and perhaps other months. One of a 
pair follows the other for about half an hour, " licking " the caudal mucus. Then 
they curve around in a semicircle with the right side of the necks in contact. In 
about fifteen minutes the lower atria are everted as small yellow eminences. These 
rapidly enlarge as eversion is completed, and the combined atria form an ovoid 
bluish-white mass between the two animals, the junction of the two atria being 
barely visible as a fine line. After three-quarters of an hour the mass shrinks and the 
animals separate, and the spermatophores are momentarily visible, partly protruding 
from the spermatheca ducts as the remainder of the shrinking mass is withdrawn. 
For further details see Quick (1946). The slender portion of the free oviduct is not 
everted as a stimulator as it is in A. hortensis, where this portion of the oviduct is 
longer. The combined mass resembles that of A. ater, since the differing situation 
of the ligula does not affect the aspect. 



BRITISH SLUGS 135 

Development 

The opaque leathery eggs, measuring from 2-25 X 2-25 to 3-25 X 2-25 mm, 
vary considerably in size and shape, even in the same cluster. Eggs are laid three 
weeks after mating. They are laid in masses of up to fifty, and tend to adhere 
together, several eggs sometimes being enclosed in a common tubular coat of mucus. 
They are white at first and become yellow or brown. Hatching occurs in from three 
to five weeks. The newly hatched young are about 6 mm long, grey, with darker 
dorsal region, and with lateral body and mantle bands. The tentacles are violet. 
Newly hatched A . ater are orange, straw colour or pale greenish-grey, and the tentacles 
dark grey rather than violet ; but the most notable difference is that the anterior 
ends of the mantle bands are closer together than in A. subfuscus (PI. 1, figs. 9 and 

11). 

Ecology 

A. subfuscus lives under logs and litter in deciduous and coniferous woods, and 
is one of the few species found in pine woods. It is also frequent in damp hedgerows, 
waste ground and some gardens. It feeds on the algal and fungal film on logs, 
and also on the larger fungi. In captivity it readily eats carrot, lettuce, oatmeal 
and dead slugs (Map 8). 

Distribution 

The whole of Great Britain and Ireland, except possibly parts of East Anglia 
and eastern Scotland. Most of Europe, north of a line through Orenburg and Odessa 
(Russia), northern Italy, Barcelona and Oporto. Introduced into North America. 

Material examined 

Living from many places in Britain. Spirit material from Britain, France, Luxem- 
bourg, Germany, Italy and Madeira. 



Arion lusitanicus Mabille, 1868 
Avion lusitanicus Mabille, 1868 : 134. La Sierra d' Arriba, near Lisbon. 

Notes 

The name lusitanicus is applied to this species with some hesitation, since Mabille 
(1868) described the tubercles as " peu allonges ". However, Pollonera's description 
and figures (1889 : 626, pi. 9) leave little doubt about the identification. A. nobrei 
Pollonera (1889) is similar to A. lusitanicus but smaller, with smaller tubercles, 
completely black, and with the distal genital ducts even more strongly pigmented. 
A specimen from Portugal in the B.M. (N.H.) collections, identified as A. nobrei, 
agrees with this description. 

Cain & Williamson (1958) discuss specimens of Arion collected at Nuneaton, 
Warwickshire, which perhaps suggest that the status of A. lusitanicus in Britain 
may require further investigation. 

ZOOL. 6, 3. 10 



136 BRITISH SLUGS 

First British record 

By Collinge (1893). Though again recorded by Collinge (1897a), the species was 
overlooked by Taylor (1902-07) and by other authors on the British fauna (e.g. 
Roebuck, 1921 ; Kennard & Woodward, 1926 ; Ellis, 1926 and 1951 ; Quick, 
1949), until its rediscovery in Durham by Mrs. Vincent (Quick, 1952). 

Diagnostic features 

Large, up to 10 cm in length, with coarse tubercles like A . ater ; from which it 
is internally distinguished by the dilated distal part of the free oviduct, which lodges 
the ligula. Distinguished from A . subfuscus by coarse tubercles and a longer sperma- 
theca duct. 

External appearance 

The length when crawling is from 7 to 10 cm. The colour varies as in A. ater, 
and may be dark grey, reddish-yellow, brown or dull greenish-grey, with darker 
head and tentacles. The transversely lineated foot fringe is sometimes darker 
and sometimes lighter than the body, and the sole is generally pale, but dark in 
some Nuneaton specimens. The respiratory orifice does not dilate to the same 
extreme degree as in A. ater. In the younger stages there is on each side a dark 
lateral band (Text-fig. 5 d). On the mantle the bands form a lyre-shaped figure, 
the right band surrounding the pulmonary orifice on all sides, and anteriorly they 
are a little closer together than in A. subfuscus of similar size. Sometimes the bands 
are retained up to maturity. 

External variation 

Var. rufescens Collinge (1893), body dark red, sides of body bandless ; var. nigres- 
cens Collinge, black or plumbaceous ; var. olivaceus Collinge, various shades of 
olive-green ; var. flavogriseus Collinge, yellowish-grey, foot fringe lighter than the 
body. 

Pallial organs, retractor muscles, nervous system 
As in A . ater. 

Jaw 

Intermediate in size (3 mm across) between those of A . subfuscus and A . ater ; 
arcuate, crossed by from nine to thirteen rather broad even ribs ; pale brown or 
yellow in light examples, nearly black in dark ones. 

Radula (Text-fig. 3 g) 

Intermediate in size between those of A. subfuscus and A. ater. The formula 
varies (in adult specimens from Durham) between C.17.35 and C.19.45. 

Alimentary system 
As in A . ater. 



BRITISH SLUGS 



137 



Reproductive system (Text-figs. 5 A and e) 

The very dark ovotestis lies a little further forward than in A. ater. The dilated 
central part of the hermaphrodite duct is folded in a zig-zag manner, and then 
narrows to form the usual " claw " at the albumen gland, which varies in size with 
the maturity of the animal. A yellowish-pink fold or ruffle accompanies the first 
part of the sperm-oviduct. The free oviduct is slender at first, and then becomes 
greatly enlarged, to narrow again slightly where it enters the upper atrium. This 
dilated part, in both the Durham and the Nuneaton examples, is deeply pigmented 
in a characteristic way (Text-fig. 5 a). The spermatheca duct is rather longer than 
in A. subfuscus, somewhat dilated at the origin. It also sometimes shows a slight 
dilatation, with stronger mucous folds, between the origin and the spherical sperma- 
theca sac. The vas deferens is about as long as the epiphallus, and the latter shows 
a conspicuous annular ring at its termination. The distal half of the epiphallus 
is deeply pigmented like the oviduct, the pigment usually tailing off in two tongues. 
In some individuals this pigmentation is less pronounced. The upper atrium is 
small, the lower pear-shaped with thick glandular-looking walls, sometimes yellow, 
and sometimes sprinkled with minute spherical white granules. 

The upper atrium contains no ligula, but the distal dilated part of the oviduct 
contains two conspicuous folds which apparently represent this structure, and 
function similarly (Text-fig. 5 e). In A. ater the ligula occupies the large upper 
atrium in both young and adult animals. Examination of the genital ducts of imma- 
ture animals of A. subfuscus and A. lusitanicus (Text-figs. 6 c and d) shows that the 
ligula lies in the oviduct, as in the adult, and does not " migrate " upwards or 
downwards. It is noteworthy that in young A . ater the oviducal branch of the genital 
retractor is inserted as in A . hortensis, A. subfuscus and A. lusitanicus ; while in 
adult A . ater (Text-fig. 6 k) the distal part of the oviduct becomes relatively much 
shorter, bringing the retractor close to the atrium. It could be argued that the 
lateral atrial pouch of A. ater is homologous with the dilated terminal portion of 
the oviduct of the other species, or alternatively that this portion of the oviduct 
is a diverticulum of the atrium. Seeing that this portion is so well devleoped in 
the early stages of growth, the second alternative is unlikely. There is in Geomalacus 
maculosus a long backward extension of the atrium ; but this carries back the 
epiphallus and the spermatheca, and does not affect the oviduct. 

Spermatophore (Text-figs. 6 h and 1) 

A chitinous tube 20 mm long, armed with a spiral row of serrations, most 
prominent towards the anterior end ; and two low smooth ridges at the posterior 
end, which die out anteriorly. 

Mating (Text-fig. 6 e) 

The figure shows the appearance of the reproductive organs just before they 
are withdrawn after mating, with the partner's spermatophore protruding from 
the spermatheca duct. Observations on the mating of A. lusitanicus are too few 
and incomplete to permit of comparison with the other species. 



138 BRITISH SLUGS 

Development 

The white eggs measure about 3 x 3-25 mm, but vary considerably in size and 
shape, both within and between clusters, and become yellower with age. In the 
laboratory, they have been laid in January and in March ; and in September by 
laboratory-bred animals only five or six months old. The clusters contain up to 
about fifty eggs, bound together by mucus, which hatch in about three weeks in 
the laboratory. 

Ecology 

In captivity, like most other slugs, A. lusitanicus is readily reared on lettuce, 
cabbage, oatmeal, etc.; but little is known of its habits in the wild. 

Distribution (Map 9) 

In the British Isles, known only from Berehaven, Bantry Bay, Ireland (Collinge, 
1893) ; the garden of No. 24, North Bailey Road, Durham and the river bank 
adjoining (Quick, 1952) ; and the site of bombed cottages at Attleborough, Nuneaton, 
Warwickshire (Cain & Williamson, 1958). On the Continent, known from near 
Lisbon, Portugal ; Mentone and Pyrenees Orientales, France ; and Basle, Switzer- 
land. The species may be indigenous to Ireland and but recently introduced to 
Durham and Nuneaton, though it seems firmly established there. Probably it is 
more widely distributed in Britain, but has not been distinguished from A . ater. 

Material examined 

Living specimens from Durham and Nuneaton. Spirit specimens from Basle, 
Switzerland, and the Pyrenees Orientales. An immature specimen from Bantry 
Bay (Collinge Collection, Museum of Zoology, Cambridge) probably belongs to this 
species. 



Arion ater ater (Linnaeus, 1758) 

Limax ater Linnaeus, 1758 : 652. Sweden. 

Arion empiricorum Ferussac, 18 19 : 60, pis. 1-3 (in part). 

First British record 
By Lister (1674). 

Diagnostic features 

Large size, coarse tubercles, wide foot fringe, large respiratory orifice ; short 
free oviduct surrounded by genital retractor muscle, ligula in upper atrium. Adults 
not certainly distinguishable in outward appearance from A. a. rufus, but usually 
black or dark brown with a grey sole, and rarely with a brightly-coloured foot 
fringe. Lateral pouch and ligula much smaller than in A . a. rufus, and vas deferens 
usually shorter. Upper atrium shorter than lower atrium. 



BRITISH SLUGS 139 

External appearance (PI. 1, fig. 3) 

Length when fully extended generally about 14 cm, but may be even longer. 
The body tubercles are long and prominent, and darker than the furrows except 
in the young. The mantle is shagreened. There is no trace of a dorsal keel at any 
stage. The caudal mucus pore is conspicuous. The sole is more or less divided 
into median and lateral areas, the lateral areas usually the darker. The wide foot 
fringe is transversely lineolated, and in dark specimens the lines extend into the 
lateral areas of the sole. In pale individuals (except true albinos, which are rare) 
the head and tentacles are much darker than the rest of the body. The respiratory 
orifice is absolutely and relatively larger than in the previously described species. 
The very sticky body mucus may be colourless, or yellowish in red and yellow forms, 
that from the caudal pore always colourless. At rest, A. ater, like A. lusitanicus, 
but unlike A. subfuscus and the smaller species, can contract to a hemispherical 
shape, or even a larger portion of a sphere. 



External variation 

Since A . a. rufus has been generally considered as only a variety of A . ater, it is 
impossible at present to tell which of the very numerous named varieties belong 
properly to the latter. Probably most of the brighter varieties belong to A. a. rufus, 
and of the duller ones to A . a. ater. 

Extensive lists of varieties are given by Taylor (1902-07), Kennard & Woodward 
(1926) and Hesse (1926), of which the following is a short selection : var. ater 
Linnaeus (1758), black with grey sole ; var. aterrima Taylor (1905), completely 
black ; var. alba Linnaeus (1758), white ; var. albida Roebuck (1883), white with 
yellow fringe ; var. castanea Dumont & Mortillet (1856), brown with paler fringe ; 
var. albolateralis Roebuck (1883) (? = var. media Jensen, 1873), dorsum black, 
sides white ; var. reticulata Roebuck (1885), tubercles pale, furrows pigmented 
(i.e. the juvenile colouring retained by the adult) ; var. hiberna Mabille (1868), 
rusty purple with paler sides ; var. plumbea Roebuck (18840), leaden grey with 
yellow fringe ; var. succinea Miiller (1774), yellow with red or orange fringe ; var. 
coccinea Gistel (1848), vermilion ; var. marginella Schrank (1803), black with red 



fringe. 



Pallial organs 

Large vessels occupy the roof and floor of the pulmonary chamber. The kidney 
is curved into a rounded quadrangular form, surrounding the pericardium, and the 
ventricle lies posteriorly to the auricle. The anterior branch of the aorta curves 
round the anterior loop of the intestine, then runs forwards supplying branches 
to the anterior part of the body, passing between the visceral and pedal ganglia. 
The posterior aorta runs backwards and supplies the alimentary and genital systems, 
and passes between the visceral and pedal ganglia. The ureter and rectum terminate 
at the posterior lip of the respiratory orifice. 



140 



BRITISH SLUGS 




Fig. 4. Avion. A. A. hortensis — internal organs in situ ( x 2) showing liver, exposed 

part of stomach, part of ovotestis, albumen gland, intestine^ aorta, crop, left salivary 
gland, nerve ring, characteristic pigmented area on buccal bulb. B. A. hortensis — 

distal genital ducts ( x 4) with position of the partner's spermatophore indicated on 
upper atrium. G. A. hortensis — distal portion of genital ducts (x 5-3) opened to 

show ligula in oviduct, glandular area below spermatheca, and papillate mucous 
membrane of epiphallus. D. A. hortensis — mating pair ( x 1-3) with slender parts 

of oviducts everted and functioning as stimulators. E. A. hortensis — individual 



BRITISH SLUGS 



141 



B 





Fig. 5. Avion. A. A. lusitanicus — reproductive organs ( x 3). B. A. subfuscus 

— distal genital ducts (x 3). C. A. ater ater — distal genital ducts (x 3). D. 

A. lusitanicus — young individual ( x 10). Compare mantle bands with those of young 
A. subfuscus and A. ater (pi. 1, figs. 9 & 11). E. A. lusitanicus — oviduct (x 5-3) 

opened to show position of ligula. 



immediately after coitus ( x 2) with everted genitalia, showing atria, everted oviduct, 
papillate epiphallus, partner's spermatophore adhering to base of spermatheca duct. 
F. A. hortensis — spermatophore ( x 6-7). G. A.fasciatus — spermatophore ( x 4), 

adapted from Lohmander. H. A. intermedius — distal genital ducts (x 4-7). 

I. A. fasciatus — distal genital ducts (x 4). K. A. hortensis — pallial region in 

ventral view, showing kidney, pericardium, auricle, ventricle, rectum and respiratory 
orifice. Muscles cut short. L. A . hortensis — cephalic and buccal retractor muscles, 

ventral view. M. A. ? hortensis — spermatophore (x 67) of a slug from a garden 
in Eaton Crescent, Swansea, Glamorgan. 



I 4 2 



BRITISH SLUGS 










Fig. 6. 



BRITISH SLUGS 143 

Retractor muscles 

The right and left cephalic retractors are widely separated at their origin, from 
the diaphragm behind the kidney. The pharyngeal retractor arises from the median 
dorsal body-wall. The genital retractor arises from the diaphragm immediately 
in front of the origin of the left cephalic retractor, ensheathes the short free oviduct, 
and sends a branch to the spermatheca duct. 

Nervous system (see Text-fig. 7 m) 

Highly concentrated in the adult, the commissures being short and the visceral 
ganglia closely united. 

Jaw 

Strong, arcuate, 3-5 mm across, crossed by a varying number of ribs (up to 
eighteen), variable in breadth, which denticulate the upper and lower edges. Colour 
yellow in pale specimens, nearly black in dark ones, the attached margin being 
darker than the free cutting edge. 

Radula (Text-fig. 3 f) 

7-8 mm long by 3 mm wide. The number of lateral teeth varies from 20 to 25, 
and of marginals from 40 to 46 on each side, but the average formula is C.22 . 44 X 130. 

Alimentary system 

The short oesophagus leads through a large wide crop to the stomach, which 
forms the posterior part of the alimentary canal. The salivary glands on each side of 
the crop are large, flattened and much lobulated. The intestine, with one anterior 
loop, is twisted through about one and a half turns — more than in the young of 
the smaller species. 

Reproductive system (Text-figs. 5 c and 6 l) 

The proximal parts resemble those of other species of Arion (see Text-fig. 5 a). 

Fig. 6. Avion. A. A. ater ater — distal genital ducts (x 10) of a young individual, 
showing long atrium and position of ligula. B. A . ater ater — genital organs ( x 3-3) 

of a nearly full-grown but immature individual, showing relative sizes of ovotestis, 
albumen gland, common duct and distal portions of ducts. C. A. hisitanicus — 

genital organs ( x 6-7) of an immature individual, showing relative sizes of organs 
and position of ligula. D. A. subfuscus — distal genital ducts (X 67) of a young 

individual, showing ligula in lower oviduct. E. A. hisitanicus — slug after coitus, 

with atrium everted. The ligula, with the oviducal orifice in its lower anterior part, 
surrounds the orifices of epiphallus and spermatheca. The partner's spermatophore 
projects from the spermatheca duct. F. A . subfuscus — distal genital ducts ( x 3-3). 
G. A. subfuscus — spermatophore (x 3-3), and portions of spermatophore (x 16-7) 
from near anterior end, centre and posterior end. H. A. hisitanicus — spermato- 

phore (x 3-5). I. A. hisitanicus — portion of posterior end of spermatophore 

(x 11). K. A. ater rufus — distal genital ducts (x 3-3). L, A. ater ater — 

distal genital ducts (x 3-3). 



i 4 4 BRITISH SLUGS 

The short free oviduct, ensheathed by the genital retractor, enters the upper atrium 
apically. It is not dilated, and does not contain the two conspicuous folds of 
A. hortensis, A. subfuscus and A. lusitanicus. However, the upper atrium contains 
a somewhat similar folded mass, arising from the walls of the atrium and the termina- 
tion of the oviduct, known as the ligula. This is much smaller than in A. a. rufus, 
where it is contained in a lateral enlargement of the atrium. The spermatheca 
duct is relatively longer than in A. intermedins, A. fasciatus, A. hortensis and 
A. subfuscus, is not bulbous at the base, and receives a branch from the genital 
retractor. The vas deferens is usually less than one and a half times as long as the 
epiphallus, which terminates in a thickened ring at the atrium. 

Spermatophore (see Text-fig. 7 l) 

This rapidly disintegrates in the spermatheca, so that it can only be obtained 
intact soon after coitus. Two specimens were obtained in this way, which are 17 
and 18 mm long. Each is somewhat dilated near the proximal end, and tapers 
in each direction. A finely serrated ridge runs nearly the whole length, the serrations 
pointing forwards. The epiphallus measures only n mm long in alcohol specimens ; 
so, unless it is much longer in life, part of the spermatophore must be formed during 
coitus. 

Mating 

Occurs in June and July, at any rate in Wales, after dark. After about half 
an hour of following and licking, the animals curve in a semicircle, neck to neck, 
and evert their atria to form a bluish- white mass. They separate after about half 
or three-quarters of an hour, when the sperm at ophores can be seen protruding from 
the spermatheca ducts, while the shrinking ligulae and atria are quickly withdrawn. 
Adams (1910), Gerhardt (1940) and Quick (1947) give fuller details. 

Development 

The eggs are variable in size, usually about 5x4 mm. They are opaque and 
leathery, pearly white when laid but turning brown. They are laid in clusters in 
the soil, two or three weeks after coitus. The first cluster may contain 150 eggs or 
more, and one or two subsequent clusters fewer than this. The eggs hatch in from 
four to six weeks. The young, 10 mm long, are pale straw colour or sometimes 
greenish, with darker head or tentacles. They are usually bandless, and if faint 
bands appear after a few days they quickly fade. Yellow pigment is concentrated 
in the tubercles. In the young of dark varieties the black pigmentation begins in 
the dorsal furrows and spreads downwards, producing a reticulate pattern which 
very occasionally persists in the adult ; though as a rule the pigment of pale adults 
is concentrated in the tubercles, leaving the furrows paler. The sole is the last 
part to pigment. 

Like the other species of Arion, A. ater shows in the later stages of embryonic 
development a cephalic and a caudal vesicle, which pulsate alternately. The tubular 
caudal vesicle persists to within a few days of hatching (Text-fig. 18 f). The jaw 
at hatching does not show any sign of bilateral origin like that of Agriolimax. 



BRITISH SLUGS 145 

Behaviour 

Adults of A. atev exhibit a peculiar rolling reflex, if touched when fully (or almost 
fully) contracted. The body sways from side to side with a simultaneous screwing 
motion, with a periodicity of about two seconds. The movement cannot always 
be elicited, but may be quite violent. As far as is known, no other slugs show this 
reflex. It is tempting to explain it in terms of alternate contraction of the cephalic 
retractors, which have separated origins in Avion. However, the reflex is only 
doubtfully elicited from A . lusitanicus, and repeated attempts to elicit it from other 
British species of Avion have always failed. 

Ecology 

A. a. atev is practically omnivorous, eating decayed vegetable matter, dead mice 
and rabbits, and animal faeces. In the lowlands, it is common in grass fields, woods, 
hedgerows and waste ground, and on grassy downs and sea cliffs. It is often the 
only mollusc present on upland peaty moors and sphagnum bogs, where it is smaller 
than in the lowlands, and either black (often the var. atevvima) or very dark brown. 

Distvibution (Map 10) 

The whole of the British Isles, northern and central Europe, Iceland and ? Russia. 
Probably some of the records from Spain, Portugal, Italy and the Mediterranean 
coasts and Atlantic isles refer to the related forms (A. a. vufus, A. lusitanicus and 
possibly others). Introduced into North America. 

Matevial examined 

Living specimens of A. a. atev, as distinct from A. a. vufus, have been identified 
from the following localities in the British Isles : Exeter, Littlehampton and 
Collompton in Devon ; Netley Heath, Surrey ; Howe Wood near Littlebury, 
Essex ; Reading, Berkshire ; Thorpe St. Andrews, Brundall Marshes and Wheatfen 
Broad in Norfolk ; Little Shelford, Homingsea and Durnford Fen in Cambridgeshire ; 
Cefn Bryn, Gower and Llangennith in Glamorgan ; Capel-y-fhn, Brecon ; Skokholm 
Island, Pembroke ; Abersoch, Caernarvon ; Bromborough, Cheshire ; Pentland 
Hills, Edinburgh ; Courtmacsherry, Cork. 



Arion ater rufus (Linnaeus, 1758) 

Limax vufus Linnaeus, 1758 : 65. ? Sweden. 
Avion empiricovum Ferussac, 1819 : 60 (in part). 
Avion sulcatus Morelet, 1845 : 28. 

Notes 

See Quick (1947). 0kland (1923) believes it to have been introduced to Sweden 
in historical times. Ferussac's empivicovum includes A. a. atev, A. a. vufus, and 
perhaps also A . lusitanicus and other species. Moquin-Tandon's description (1855-56, 
2 : 10) of empivicovum, and his fig. 1 on pi. 1, could serve for either A. a. atev or 



146 BRITISH SLUGS 

A. a. rufus ; but his fig. 12 represents the genitalia of A. subfuscus, A. lusitanicus 
or a similar species. 

Cain & Williamson (1958) have studied pigmentation and genital anatomy in 
populations referable to A. a. ater and A. a. rufus. Their evidence suggests that only 
the former is indigenous to the British Isles, where A. a. rufus is an introduced 
alien associated with cultivated land. In Europe the natural ranges of the two forms 
seem not to overlap widely, since A. a. ater apparently occupies Scandinavia and 
A. a. rufus much of the rest of the continent, but it is not yet known how the two 
forms interact along their natural boundary. However, two populations near 
Oxford show the intermediacy of character combined with high variability which is 
to be expected from hybrids. This suggests that the two forms should be considered 
as geographical races. 

First British record 

By Gerhardt (1940), on specimens from Swansea, Glamorgan. The subspecies 
had been considered by previous British authors as merely a variety of A . a. ater. 

Diagnostic features 

Adult not certainly distinguishable from A. a. ater on external appearance, but 
usually yellow to reddish-brown, with a creamy yellow sole and bright orange or 
red foot fringe. Internally, upper atrium longer than lower, much larger ligula in 
lateral pouch, vas deferens usually longer than in A. a. ater. Young have mantle 
bands, rarely found in A. a. ater. 

External appearance, external variation, pallial organs, retractor muscles, nervous 
system, jaw, radula, alimentary system 
As in A . a. ater. 

Reproductive system (Text-fig. 6 k) 

As in A. a. ater, except for the distal genital ducts. The upper atrium is larger, 
and exhibits a lateral bulging that lodges a much larger ligula. The lower atrium 
is shorter than the upper. The vas deferens is relatively longer, usually more than 
one and a half times as long as the epiphallus and sometimes nearly twice as long. 
The oviduct, instead of entering the apex of the atrium formed by the lateral bulge, 
enters between this and the entrance of the epiphallus. Sometimes (as in specimens 
from Limeslade Bay, Swansea) the lateral bulge is narrower and prolonged further 
backwards. Occasional specimens of A . a. ater show a suggestion of a lateral bulge, 
but the size of the ligula, and the relative sizes of upper and lower atria and the vas 
deferens and epiphallus, serve to identify it. 

Spermatophore (Text -fig. 7 l) 

Probably longer than that of A. a. ater (about 25 mm long), and even more 
disproportionate to the size of the epiphallus in spirit material. This may be associ- 
ated with the longer period of coitus, if the spermatophore is formed during coitus. 



BRITISH SLUGS 147 

Mating 

A. a. rufus attains maturity later in the year than A . a. atev, and mates in September 
and October, at any rate in South Wales. Mating resembles that of A. a. ater, 
except that coitus lasts for two hours. For further details, see Gerhardt (1940) 
and Quick (1947). 

Development 

The eggs are like those of A. a. ater. The newly-hatched young are pale yellow 
or orange, and differ from those of A. a. ater in having (at birth or after a few days) 
dark mantle and body bands, which persist for a few months. In individuals which 
will be dark when adult, the course of pigmentation is as in A. a. ater. The young 
can be distinguished from those of A. subfuscus by their coarser tubercles, and by the 
closer approximation anteriorly of their mantle bands (PL 1, figs. 9 and 11). 

Behaviour 

A. a. rufus shows the same rolling reflex as A. a. ater. 

Ecology 

In Britain, A. a. rufus is more likely to be found in gardens and parks (though 
sometimes at a considerable distance from dwellings) than in wilder places. It 
never seems to occur in peaty moors or sphagnum bogs ; though this may reflect 
its introduction by man, and consequent restriction to cultivated land, rather than 
dependence on calcium in the soil. 

Distribution (Map 11) 

Not fully known, since A. a. rufus has only recently been distinguished sub- 
specifically from A. ater. Authentically known in Britain from Glamorgan, Somerset, 
Berkshire, Oxford, Essex, Cambridge and Leicestershire. It seems to be the native 
form in Europe south of Denmark, though the natural ranges of the two subspecies 
are insufficiently known. 

Material examined 

Living specimens from Monkton Coombe near Bath, Somerset ; Chingford, 
Essex ; Basildon, Berks ; Coombe End, Oxford ; Lords Meadow near Cambridge ; 
and the Swansea district, Glamorgan. Spirit material from Loughborough, Leicester- 
shire ; Borgholm, Oland Island, Sweden ; Froedensborg, NE. Zealand, Denmark ; 
Canton Solothurn, Switzerland ; and from France, Corsica and Portugal. 



Superfamily Zonitacea 

Family Limacidae 

Slugs with a small calcareous shell almost always completely enclosed by the 
mantle. The dorsum is keeled posteriorly or right up to the mantle. There is no 



BRITISH SLUGS 



caudal mucus gland. The sole is tripartite and a supra-pedal groove is present 
(aulocopod). The jaw is smooth, usually with a median projection (oxygnath), 
and the lateral radular teeth are tri- or bicuspid, the marginal teeth aculeate. The 
spermatheca duct is short, generally arising from the atrium. 




tzz^iSimWk 


'T-rft^Tf 


c 




^K^« 


IH?^ 


\ — \ 




<^> 




Wa. 



Fig. 7. Limacidae. External appearance. A. Milax — keel extending forward to 

mantle ; mantle granulate with horseshoe furrow ; respiratory orifice behind centre 
of mantle ; genital orifice between upper tentacle and respiratory orifice. B. 

Agriolimax — keel extending only a little way forward ; tail truncate ; mantle con- 
centrically ridged with nucleus over respiratory orifice, behind centre of mantle ; genital 
orifice close behind upper tentacle. G. Limax — keel extending a little way forward ; 

tail not truncate ; mantle concentrically ridged with nucleus in median dorsal line ; 
respiratory orifice behind centre of mantle ; genital orifice close behind upper tentacle. 
D. Milax — chevroned foot-sole, stimulator protruded. 

E. Lehmannia marginata — jaw. F. Agriolimax veticulatus — jaw of very young 

individual, showing bilateral origin. G. Avion hovtensis — jaw, for comparison. H. 

Limax maximus — shell. 

Avion ater ruf us. I. Pair in copula. K. Portion of posterior end of sperma- 

tophore (x 24). L. Spermatophore (x 1-7). M. Ganglia and commissures 

of nerve ring (buccal ganglia omitted). 



BRITISH SLUGS 149 

Subfamily Parmacellinae 

The shell is paucispiral, not completely enclosed by the mantle (in Parmacella) , 
or flat and enclosed (in Milax) . The mantle is granular, bearing a horseshoe-shaped 
groove, and the respiratory orifice lies behind the middle of the right margin. The 
genital orifice is between the respiratory orifice and right upper tentacles. The 
dorsum is keeled up to the mantle. The central and lateral radular teeth are tricuspid. 
The intestine has one forwardly directed loop, and the rectum lacks diverticula. 
An epiphallus secretes a spermatophore, and atrial glands are present. Milax is 
the only British genus. 

Watson (1930) considers that Parmacella and Milax arose from a common stock ; 
Parmacella, with its spiral shell, embedded pedal gland, and ocular retractor crossing 
the penis, being the more primitive. 



Genus MILAX Gray, 1855 

Type species Limax gagates Draparnaud, 1801. 

The shell, completely enclosed by the mantle, has a median non-spiral nucleus 
near the posterior margin. The pedal mucus gland lies free in the body cavity, 
and not embedded in the foot below it, as in the other genera. The right ocular 
retractor lies to the left of both penis and vagina. In M. sowerbyi and M. gagates 
an atrial stimulator is present. The median area of the sole is crossed by A-shaped 
grooves. Milax is without lateral body bands at all stages of growth. 

Watson (1930) considers that, of the British species of Milax, M. gagates (with 
more lateral and fewer marginal radular teeth, and a wider distribution) is the most 
primitive, and M. budapestensis (with fewer lateral and more marginal teeth, and 
no stimulator) the most specialized. 

A common continental species, M. marginatus (Draparnaud, 1801) (= M. rusticus 
Millet, 1843), which it seems possible may yet be discovered in Britain, is a yellowish 
slug with small black spots on the body and mantle and a dark band on each side 
of the latter, and resembles M. budapestensis in lacking the atrial stimulator. 



Milax gagates (Draparnaud, 1801) 

Limax gagates Draparnaud, 1801 : 100. Presumably near Montpellier, France. 
Amalia parryi Collinge, 1895a : 7. 
Amalia babori Collinge, 1897c : 294. 

Notes 

Two species are currently confused under the name gagates. The common British 
species is distinguished by its smooth genital stimulator, and less certainly by its 
smoother appearance (with flattened tubercles) and paler sole. The second species 
is primarily Mediterranean in distribution, though there is one hitherto unpublished 



i5o BRITISH SLUGS 

British record (p. 156). It has two rows of large papillae on the stimulator, and also 
more prominent tubercles and a darker sole. 

It is not easy to determine the proper application of the name gagates. There 
are no slugs among Draparnaud's types at the Naturhistorisches Museum, Vienna 
(Locard, 1895 : 154; confirmed by Dr. O. E. Paget in litt.). Both species occur 
at Marseilles, only eighty miles from Draparnaud's home at Montpellier. Drapar- 
naud's description does not deal with the internal anatomy. However, he describes 
gagates as " tres-lisse, tres-luisante " (1801) and " nitidus ; corpore striato subrugoso" 
(1805). This strongly suggests the north-western species, since he applies " sub- 
rugosus " to other slugs as smooth as this, and uses " rugosus " only for those as 
rough as the Mediterranean species. 

Few early authors described or figured the condition of the stimulator, from which 
alone their application of the name gagates could be determined. Lessona & Pollonera 
(1884 : 105, pi. 2, fig. 2) use this name for Italian slugs with papillate stimulators. 
Germain (1930, fig. 52 a), in treating the French fauna, gives a figure of a papillate 
stimulator which is apparently copied from Lessona & Pollonera, and does not 
mention the locality. Taylor (1902-07, fig. 159) shows the smooth stimulator of a 
British slug. 

The name parryi Collinge is a synonym of gagates, as the latter is here interpreted, 
since one of the type specimens from Santa Cruz, Teneriffe, shows on dissection a 
penis, epiphallus and stimulator like the British species. So does one of a series from 
Haleakala Maui, Hawaii, apparently the types of babori Collinge. The following 
names, among others, cannot at present be allocated to either of the species with any 
certainty : 

Limax maurus Quoy & Gaimard, 1824 : 4 2 7- P° r t Jackson, Sydney, Australia. 

Parmacella nigricans Schultz, 1836, 1 : 125. Palermo, Sicily. 

Amalia mediterranea Cockerell, 1891 : 331. Algeria. 

Amalia mediterranea forma similis Cockerell, 1891 : 332. Catania, Sicily. 

However, specimens that I received from Catania in 1947 proved to be the species 
with a papillate stimulator. 

First British record 

By Thompson (1840 : 205). 

Diagnostic features 

Uniform grey or black colour, with darker keel, respiratory orifice without pale 
margin, mucus colourless ; smooth genital stimulator, rather thick spermatophore, 
not hooked at the thick end and with spines distributed more evenly than in M. 
sowerbyi. 

Differs from M. cf . insularis in smooth, not papillate, stimulator ; and less certainly 
in smoother skin and paler sole. 

External appearance (PL 1, fig. 12) 

Length extended about 5 cm. When contracted, the slug is more compressed 
laterally than M. sowerbyi. The colour is usually grey or black, getting lighter 



BRITISH SLUGS 151 

towards the foot. The tubercles are flattened, making the slug rather smooth, 
and the intervening grooves are unpigmented. The keel is usually dark, and is 
abruptly angulated at the posterior end. The central area of the granulate mantle 
is delimited by a horseshoe-shaped groove, open posteriorly, with the arms angulated 
at the level of the respiratory orifice. The respiratory orifice is small and without 
a pale border, and is situated a little behind the centre of the right margin of the 
mantle. The sole is pale, distinctly tripartite, with the median zone crossed by 
A-shaped grooves (Text-fig. 7 d). Only this central area shows the locomotory 
waves. The mucus is sticky and colourless. 

External variation 

Var. plumbea Moquin-Tandon (1855), paler or darker grey; var. rava Williams 
(1888), drab coloured, slightly fuscous, mantle paler ; var. bicolor Taylor (1904), 
deep red on the sides ; var. benoiti Lessona & Pollonera (1882), black, keel whitish. 

Other varieties with a Mediterranean distribution mentioned by Taylor (1902-07 : 
143-146) probably refer to M. cf. insularis. 

Shell (Text-fig. 8 k) 

Measures about 4 X 2-5 mm. White and shining. Somewhat saddle-shaped 
above, and flattened below. The nucleus is near the posterior margin, in the middle 
line, and the lines of growth are distinct. Viewed laterally, the shell is concave 
from back to front, with the nucleus at the highest point, and shows a groove separat- 
ing the upper part of the shell from the lower. 

Pallial organs 

The kidney has an extension to the right toward the rectum, and the auricle 
and ventricle lie anterior to the left half of the kidney. The rectum crosses over the 
common origin of the tentacular and pharyngeal muscles to the posterior lip of 
the respiratory orifice. 

Pedal gland 

Lies free in the body cavity, occupying half the body length. Its duct projects 
posteriorly from the glandular tissue for about 0-25 mm. 

Retractor muscles 

The pharyngeal and tentacular retractors are not separated, as in Arion, but have 
a common origin from the posterior edge of the diaphragm in the middle line. At 
the posterior border of the kidney this divides into the pharyngeal retractor, which 
is bifurcate for about one-third of its length, and the tentacular retractor. The latter 
soon divides into right and left branches, and these divide again much further 
forward into branches to the upper and lower tentacles. The penial retractor arises 
further forward, on the left side of the diaphragm. 

ZOOL. 6, 3. 11 



152 



BRITISH SLUGS 




«o 













j/H 



J/ 



Fig. 8. Milax. Distal genital ducts ( X 4). A. M. gagates — atrium opened to show 
stimulator. B. M. sowerbyi — atrium opened to show stimulator. C. M. buda- 

pestensis. D. M. cf. insularis. 

Spermatophore (x 6-7), spines (x 33). E. M. gagates — one spine. F. 

M. sowerbyi — two spines. G. M. budapestensis — three spines. 

H. M. cf. insularis — papillate stimulator. 

Shell ( x 2-7), dorsal and lateral views. I. M. sowerbyi. K. M. gagates. 

L. M. budapestensis. 



BRITISH SLUGS 153 

Nervous system 

The cerebral ganglia are connected by a short broad commissure. The cerebro- 
pleural and cerebro-pedal commissures are short, so that the nerve ring closely 
surrounds the oesophagus. The pleural, visceral and abdominal ganglia are closely 
approximated, as are also the visceral and pedal ganglia. The abdominal ganglion 
is almost completely fused with the left visceral. There is little if any variation in 
the nervous system between the species of Milax. 

Jaw 

Brown, oxygnathous, measuring 17 mm across. 

Radula (Text-fig. 9 a) 

The central tooth has a mesocone and well-developed ectocones. The laterals 
have well-marked endo- and ectocones, nearly equal in size. Most of the marginals 
lose the ectocones, but five or six of the innermost teeth retain them. Formula 
C.17.27, with slight variation (see p. 157). 

Alimentary system 

The brown oesophagus leads to a long capacious crop, and the intestine (which 
exhibits marked spiral torsion) extends well behind the stomach, thus contrasting 
strongly with the condition in Arion. The rectum runs forwards above the common 
stem of the retractor muscles. 

Reproductive system (Text-fig. 8 a) 

The spiral ovotestis, mainly hidden by the upper lobes of the liver, gives off the 
pale hermaphrodite duct. This is much folded in its anterior wider part, and the 
terminal slender portion bears an oval seminal vesicle at its entry into the linguiform 
albumen gland. The common duct is long, voluminous and folded. The spermatheca 
is an oval sac, with a short duct arising from the oviduct 1-5 mm before its termina- 
tion. 

The prostate is a yellow ribbon-like organ on the common duct. At the beginning 
of the free oviduct it gives off the somewhat convoluted vas deferens, which enters 
the apex of the epiphallus. The epiphallus is widest apically and has a rounded 
eminence or blunt caecum below the apex, a feature that is not shared by M. sowerbyi 
or M. budapestensis. At the insertion of the retractor muscle, the epiphallus, without 
any external sign, passes into the penis, which narrows towards its termination in 
the atrium. Before its termination the penis shows two more or less distinct dilata- 
tions, the upper one containing the blunt perforate penial papilla with a small 
pigmented diverticulum. The mucous membrane of the epiphallus is thrown into 
transverse circular folds, and that of the penis into longitudinal folds. A lobulated 
gland that can be unravelled into long glistening white tubules opens into the atrium. 
The atrium lodges a fleshy conical stimulator attached by its larger end, with the 
tip often folded back. The stimulator is generally smooth but occasionally has two 
or three minute papillae near the tip. 



154 



BRITISH SLUGS 






n '<* 30 ^ 





c » 



7 n »« 



B 











30 1+0 




17 iq 30 uo 







^ 



» 2. 3 








C I 




SSow 



c 1 * 



3 5 




|l* tK row 



3 7 

9 p 



20*^ row 



7 '0 




i? e 



Fig. 9. Representative radular teeth ( x 330). A. M. gagates. 



It-0 row 



B. M. sowerbyi. 



C. M.budapestensts. Representative radular teeth 

M. sowerbyi, an individual one day old. D. Represent axive 

(X 1,000). E. Jaw (X 67). 



BRITISH SLUGS 155 

Spermatophore (Text-fig. 8 e) 

A brown chitinous tube about 10 mm long, widest at the centre and narrowing 
at the anterior end. Except at two places on the concave side, it bears prominent 
spines throughout. The spines have four main branches, and fourteen or sixteen 
terminal points. 

Mating 

Observations are very scanty, but apparently the partners during coitus are in 
close contact, and little or nothing of the genitalia is visible. Though the stimulator 
is often found protuded in spirit specimens, it is not visible during coitus. Mating 
has been recorded in spring, summer and winter. Taylor describes the eggs as 2 X 1*5 
mm, transparent and thin shelled. This, if correct, is much smaller than the eggs of 
M. sowerbyi and M. budapestensis. 

Ecology 

Though widely distributed in the British Isles, M. gagates seems to be commoner 
near the sea than inland, and especially common in the south-west. In parts of 
Devon it is abundant in root crops, and can be a pest. It inhabits cliffs, wild places 
and gardens. In captivity, like most other slugs, it eats carrot, potato and oatmeal. 

Distribution (Map 12) 

M. gagates probably occurs in all the vice-counties of England, Wales and Ireland, 
the south of Scotland, the Isle of Man and the Clyde Islands. In Europe, its distribu- 
tion is essentially north-western, though specimens from Marseilles have been 
examined. Other records from the Mediterranean area probably refer to a closely- 
related species, here distinguished as M. cf. insularis (p. 156). 

M. gagates has been introduced into South Australia, Hawaii, California, Canary 
Isles and Tristan d'Acunha. It is not yet clear whether other records of introduc- 
tions into America, Bermuda, Ascension Island, St. Helena, South Africa, Australia, 
Tasmania, New Zealand and Polynesia refer to M. gagates or to M. cf. insularis. 

Fossil record 

Pleistocene to Recent. 



Material examined 

Living specimens from Epsom, Surrey ; Southwell, Nottingham ; Porthcawl, 
Glamorgan ; Haverfordwest, Pembrokeshire ; St. Agnes, Scilly Isles ; and Majorca. 
Spirit specimens from Aberdeen, and from Marseilles, France ; Santa Cruz, Teneriffe ; 
Calbaden Canyon, Puenta Hills, Los Angeles, California ; Kensington suburb, 
Adelaide, South Australia (coll. B. C. Cotton) ; and Haleakala Maui, Hawaii. 



156 BRITISH SLUGS 

Milax cf. insularis (Lessona & Pollonera, 1882) 

Amalia insularis Lessona & Pollonera, 1882 : 57 (see 1884 : 103). Sardinia and Sicily. 
Amalia ichnusae Lessona & Pollonera, 1882 : 60 (see 1884 : 106). Sardinia. 

Notes 

If the name gagates is to be reserved for the north-western species with the smooth 
stimulator (p. 149), the valid name for the present species remains to be determined. 
Several names whose application is uncertain (some of which are listed on p. 150) 
are available within the gagates complex. It will require a thorough study of the 
Mediterranean forms to clarify the biological and nomenclatural situation. 

First British record 

This is the first record to be published. Mr. Armitage collected specimens near 
Bexhill, Sussex, about 1948, and the differences between these and M. gagates 
were confirmed by Mr. Watson {in litt.). All other specimens examined, from various 
parts of Britain, have proved to be M. gagates. 

Diagnostic features 

Differs from M. gagates in having large papillae on the atrial stimulator. Also, 
the slug is usually larger, blacker and less smooth, with a darker sole. 

External appearance 

As M. gagates, except for the usually larger size, more prominent tubercles, and 
more intensely black coloration, with the sole not conspicuously paler. 

External variation 

It is not at present possible to assign named varieties to this poorly understood 
species. 

Shell, pallial organs, pedal gland, retractor muscles, nervous system 
As in M . gagates, as far as is known. 

Jaw 

As in M. gagates, though varying from pale brown to nearly black in the material 
examined. 

Radula 

The teeth closely resemble those of M. gagates in form and number. Formulae of 
radulae examined were : M. cf. insularis C.18.28 (Catania, Sicily), C.17.28 x 105 



BRITISH SLUGS 157 

and C.19.27 (Malta), C.20.30 X 102 (Marseilles) ; M. gagates C.15.31 X 100 
(Aberdeen), C.16.27 (Porthcawl), C.17.27 X 96 (California), C.19.27 X 102 
(Marseilles). 

Alimentary system 
As in M. gagates. 

Reproductive system (Text-fig. 8 d) 

Here the differences from M. gagates are obvious. The stimulator (Text-fig. 
8 h), instead of being smooth or having at the most two or three minute papillae 
near the tip, has large papillae disposed in four longitudinal rows near the base, 
diminishing to two rows and becoming smaller towards the apex. Sometimes the 
papillae are mucronate, and in one specimen they were found to be partly retracted. 
The atrial glands extend further backwards than in M. gagates, in a tongue-like 
prolongation. 

The epiphallus does not possess the sub-apical bulge seen in M. gagates. The small 
internally-pigmented accessory penial papilla, instead of being enclosed in the 
penis sheath, bulges externally ; but it seems likely that this character varies with 
the physiological state of the animal, in both species. 

Whereas Mediterranean specimens examined had four rows of papillae towards 
the bases of their stimulators, Lessona & Pollonera (1884, pi. 2, figs. 1 and 2 ; 6 and 
7) figure "gagates " and insularis with only two rows. They describe ichnusae as 
having a single row of papillae, though their figure shows the stimulator as smooth 
(1884 : 106 and pi. 2, fig. 4). Possibly there is more than one species with a papillate 
stimulator in the Mediterranean region. Mr. Aten of Zaandam, Holland, reports 
(in litt.) finding M. gagates and M. cf. insularis in the Pyrenees Orientales, and that 
the latter when young has only a single row of papillae. This suggests that M. ichnusae 
may be an immature form. 

Spermatophore, mating, development, behaviour, ecology 
Not known. 

Distribution 

In Britain, once recorded from Bexhill, Sussex. In Europe, found in the Mediter- 
ranean region. Occurs together with M. gagates near Marseilles. 

Fossil record 

Not distinguishable from M. gagates. 

Specimens examined 

Spirit specimens from Marseilles and Valence, Malta, Sicily and Majorca. 



158 BRITISH SLUGS 

Milax sowerbyi (Ferussac, 1823) 

Limax sowerbyi Ferussac, 1823, 2 : 96, pi. 8. London. 

Limax carinatus Risso, 1826 : 56. 

Limax marginatus Jefferies, 1862 : 132. 

Amalia maculata Collinge, 18956 : 336, pi. 23, fig. 6. 

Amalia collingei Hesse, 1926 : 139. 

First British record 
By Ferussac (1823). 

Diagnostic features 

Differs from M. gagates in brown colour speckled with darker patches, pale keel 
and margin of respiratory orifice, grooves between tubercles pigmented, keel not 
abruptly truncated, animal less compressed laterally when contracted, mucus yellow 
and more tenacious. Differs from M. budapestensis in broader, uniformly pale sole. 

Internally characterized by short blunt atrial stimulator, long conical spermatheca, 
tapered spermatophore hooked at blunt end and bare of spines at narrow end. 

External appearance (PI. 2, fig. 23) 

Length when extended about 7 cm. When contracted the slug is rather less 
compressed laterally than M. gagates, and the keel is not abruptly truncated behind. 
The body is usually pale or dark brown with darker speckling, with the keel paler 
and the grooves between the tubercles pigmented. The mantle grooves are as in 
M. gagates, but the respiratory orifice is pale margined. The tripartite sole is pale, 
and the sticky mucus yellow. 

Variation 

Var. alba Taylor (1904), entirely white ; var. fuscocarinata Cockerell (1886c), 
keel the same colour as the back ; var. nigrescens Cockerell, black with orange 
keel ; var. oretea Lessona & Pollonera (1882), with median dark streak on mantle 
— found in Devon and Glamorgan. 

Shell (Text-fig. 8 1) 

Larger, thicker and more evenly oval than that of M. gagates, measuring 5x3 mm. 

Pallial organs 
As in M. gagates. 

Pedal gland 

The duct does not project beyond the glandular tissue. 

Retractor muscles 

The pharyngeal retractor is more deeply cleft than in M. gagates, and the two 
tentacular retractors separate sooner. The penial retractor arises nearer to the 






BRITISH SLUGS 159 

common stem of the other muscles, and there is a pair of short muscles from the 
penis to the body-wall. 

Nervous system 
As in M. gagates. 

Jaw 

As in M. gagates, but a little broader from upper to lower margin. 

Radula (Text -fig. 9 b) 

As in M. gagates, except that there are a few more transverse rows of teeth, and 
there are fewer laterals and more marginals in each row, with two or three of the 
innermost or transitional marginals often retaining a minute ectocone. Formula 
C.13.36 X 115. 

Alimentary system 
As in M. gagates. 

Reproductive system (Text-fig. 8 b) 

The lower genital ducts differ from those of M. gagates in several features. The 
short wide spermatheca duct arises from the free oviduct even nearer the atrium, 
and the sac has a narrowing backward prolongation. The epiphallus widens instead 
of narrowing towards its termination, and does not present the sub-apical lateral 
bulge of M. gagates, nor the small pigmented diverticulum at the distal end of the 
penis. The atrium contains a short blunt smooth stimulator. 

Spermatophore (Text-fig. 8 f) 

Thicker than in M. gagates, measures about 13 mm long. The thinner anterior 
end is smooth, and the curved posterior end bears recurved spines of three types. As 
usually found in the spermatheca, and figured by Taylor (1902-07, fig. 174) and 
others, the smooth anterior end terminates abruptly. However, if the spermatophore 
is taken very shortly after coitus, before the spermathecal fluid has had time to act 
upon it, the anterior end is found to be capped by a curious conical structure about 
2 mm long, with a rounded apex, and formed by four leaflets with fringed edges 
(Quick, 1950). A fresh spermatophore of M. budapestensis does not have this cap, 
but no opportunity has occurred of examining a fresh spermatophore from M. 
gagates. Occasionally two spermatophores are found in the spermatheca in M. sowerbyi, 
but whether they are transferred during a single coitus (which is very prolonged) 
or on two separate occasions is not known. 

Mating 

After following and then curving around, the pair remain in close contact for 
twelve to eighteen hours, and little or nothing is visible of the genitalia. Pairing 
has been observed in October and November, but perhaps occurs at other times. 



160 BRITISH SLUGS 

Development 

The eggs are 4x3-5 mm, soft, translucent and pale amber, laid in clusters of 
about a dozen, hatching in from four to six weeks. The embryo before hatching 
has a large flattened caudal vesicle nearly circular in outline. When hatched the 
young are 8 mm long, with a conspicuous pale keel. The mantle is speckled with 
black and sometimes has a dark median streak, which is retained in var. oretea when 
adult. 

Ecology 

M. sowerbyi is found in gardens, cultivated fields and rubbish heaps. It burrows 
in the ground, and can do considerable damage to carrot and potato crops. 

Distribution (Map 13) 

Occurs in probably every vice-county of England, Wales, Ireland and the southern 
half of Scotland, and in France, Belgium and Spain. Probably known under other 
names from Portugal, Italy, Greece and the north coast of Africa. Introduced into 
South America and New Zealand? 

Fossil record 

Pleistocene to Recent. 

Material examined 

Living specimens from many places in Glamorgan, Pembroke, Surrey, Berkshire 
and Sussex. Spirit material from Britain and Marseilles. 



Milax budapestensis (Hazay, 1881) 

Limax gracilis Leydig, 1876 : 276, nee Limax gracilis Rafinesque-Schmaltz, 1820a. 
Amalia budapestensis Hazay, 1881 : 40. Budapest. 

First British record 

By Phillips & Watson (1930). 

Diagnostic features 

Sole with dark central and paler lateral areas ; body when contracted tends to 
lie in a curve and is less compact and humped than M. gagates and M. sowerbyi. 
No atrial stimulator, spermatophore slender and spirally twisted. Radula has 
more marginal teeth, and spermathecal duct is longer, than in M. gagates and 
M. sowerbyi. 

External appearance (PL 2, fig. 20) 

The smallest British species of Milax, very slender when extended to 5 cm. 
The dorsum is dark, sometimes nearly black or with a brownish tinge, with a dirty 



BRITISH SLUGS 161 

yellow or orange keel. The grooves between the dorsal and the lateral, somewhat 
polygonal, tubercles are deeply pigmented. The mantle is sometimes paler than the 
dorsum, the horseshoe-shaped groove deeply pigmented, and the small respiratory 
orifice has a grey margin. The peripodial groove and margin of the sole are deeply 
pigmented, the lateral areas of the sole grey, and the median area dark grey or nearly 
black. The mucus is viscid and colourless, but in the mantle area becomes yellowish 
after repeated irritation. 

External variation 

Variation is confined to the intensity of the general grey colour ; to the tint of 
the keel, which varies from yellow to brownish-orange ; and to the sole, whose 
lateral areas are occasionally darker than the central area. 

Shell (Text-fig. 8 l) 

Concave or flat below, measuring 3 X 1-25 mm, brownish in colour. 

Pallial organs 
As in M. gagates. 

Pedal gland 

The duct extends behind the glandular tissue for about 5 mm. 

Retractor muscles 

The tentacular and pharyngeal retractors are usually rather more deeply cleft 
than in M. gagates, less so than in M. sowerbyi. 

Nervous system 
As in M. gagates. 

Jaw 

Brown and oxygnathous as in the other species but smaller, measuring about 
1*5 mm across and 0-5 mm in height. 

Radula 

Differs from those of the other species in having the basal plate of the central 
tooth relatively broader, the aculeate mesocones of the marginals more slender, 
fewer lateral teeth and more marginals. Formula C. 7-8. 38-40. 

Alimentary system 
As in M. gagates. 

Reproductive system (Text-fig. 8 c) 

The pale spirally twisted ovotestis has five or six closely approximated lobes, 



162 BRITISH SLUGS 

each composed of numerous follicles. The hermaphrodite duct is slender for the 
greater part of its length, but becomes dilated and folded near the albumen gland, 
and here shows a small seminal vesicle. The common duct with a broad yellow 
prostatic ribbon is folded S-wise, and after parting from the vas deferens contracts 
to form a relatively long slender free oviduct. At about two-thirds of its length, 
the wider relatively long spermatheca duct arises, leading to a cylindrical sac with 
a rounded apex. The rather long flexuous vas deferens enters the epiphallus sub- 
apically. In the writer's experience the epiphallus and penis are long and slender, 
but Phillips & Watson (1930) describe and figure a short blunt form in addition, and 
find this the commoner. It is not known if this is a true dimorphism, or only the 
result of the physiological condition. Hazay (1881, pi. 1) shows an intermediate 
form of penis and epiphallus, and Simroth (1885, pi. 10, fig. 13) the long form. Both 
forms occur in Belgian specimens, sent to me by Dr. W. Adam. The atrium receives 
the slender ducts of a pair of lobulated atrial glands, and has no stimulator. 

Spermatophore (Text-fig. 8 g) 

Length about 16 mm, anterior end slender, posterior end spirally twisted. No 
trace of a frilled cap as in M. sowerbyi, even in sperm at ophores removed during or 
immediately after coitus. Except for the first 2 mm at the anterior end, the spermato- 
phore is covered by forwardly-recurved branching spines, which are smaller and 
less branched towards the ends. There are two rows of spines anteriorly, three or 
four rows on the posterior two-thirds. 

Mating 

Occurs during November, December and January, under cover of loose stones 
or logs. Coitus generally starts in the evening and continues until the following 
mid-day or later. A little of the partly everted atria and the two penes is visible 
between the necks of the partners. The atrial glands are partly contained in the 
everted atria, together with the distal ends of the genital ducts. 

Development 

The ellipsoidal yellow leathery eggs are laid in small clusters underground, and 
measure 2-9 X 2-25 mm. Those laid in December hatch in March. The young 
at hatching are 4-5 mm long, pale yellowish-grey with a recognizable keel. 

Behaviour 

Whereas when at rest the other species of Milax assume a compressed helmet 
shape, M. budapestensis is usually found only half contracted, and curved into a 
semicircle. 

Ecology 

Occurs mostly in gardens and allotments, under stones and rubbish, and can do 
considerable damage to roots and tubers. Though much less common in wild places, 
there are records for such situations in Wales and Cheshire. 



BRITISH SLUGS 163 

Distribution (Map 14) 

Recorded from many localities in the British Isles, mainly in western England and 
Wales, but also in south-eastern England, southern Ireland and the Hebrides. 
On the Continent, known from Belgium, Germany (Wurtemberg, Tubingen, the 
Rhon Mountains and Wiirzburg), North Italy (Verona, Vicenza and Padua), 
Hungary (Budapest), Transylvania, Bulgaria and Crimea. It probably occurs in 
many places between these widely-separated localities, in the zone from the British 
Isles to the Crimea. For a map and further details, see Phillips & Watson (1930) 
and Ellis (1951). 

Fossil record 
Recent. 

Material examined 

Living specimens from Berkshire, Glamorgan, Pembroke and Denbighshire. 
Spirit material from Britain and Belgium. 



Subfamily Limacinae 

Limacidae in which the keel rarely extends forwards as far as the mantle. The 
nucleus of the enclosed shell is terminal and lies to the left of the mid-line. The mantle 
is concentrically ridged and lacks a horseshoe-shaped furrow. There is no well- 
developed epiphallus, no spermatophore and no atrial gland. The stimulatory 
organ, when present, is a penial sarcobelum, and not an atrial stimulator as in 
Milax. The endocones of the lateral radular teeth, when present, are more or less 
united with the mesocones. The intestine has one or two forwardly-directed loops. 



Genus AGRIOLIMAX Morch, 1865 

Type species Limax agrestis Linnaeus, 1758. 

The right ocular retractor lies to the left of both penis and vagina. The intestine 
has one forwardly-directed loop, and the right lobe of the liver forms the apex of the 
visceral mass. A penial sarcobelum is present, more or less developed. The nucleus 
of the concentric mantle ridges lies to the right of the middle line, over the respiratory 
orifice. The tail is obliquely truncate. 

Pilsbry (1944, 1948) adopts the name Deroceras Rafinesque-Schmaltz (1820^ : 10) 
for this genus, as do most American and many European authors. Watson 
(1943 : 54) discusses the question. Ellis (in press) has submitted to the International 
Commission on Zoological Nomenclature a proposal for the suppression of Deroceras 
and other older names, and the validation of Agriolimax. 

Hesse (1926) mentions sectional names for the first three species considered here. 
Agriolimax s.s. (including A. agrestis and A. reticulatus) has a rectal caecum and 



i6 4 BRITISH SLUGS 

a more or less tufted penial appendix, whereas Hydrolimax Malm (including A . laevis) 
lacks both caecum and appendix, but has a long tubular penial diverticulum. Other 
sectional names have been proposed for groups of extra-limital species, such as 
Krynickillus Kaleniczenco, which lacks caecum, appendix and sarcobelum. However, 
several species show combinations of characters not covered by existing sectional 
names. A. caruanae has a tufted penial appendix like Agriolimax s.s., but lacks a 
rectal caecum like Hydrolimax ; while one or both of its penial diverticula may 
correspond to the latter's single but minutely bifid diverticulum. A slug collected 
in Cyprus by A. R. Waterston (? A. cyprius Simroth, 1906, or A. panormitanus 
Lessona & Pollonera, 1882) has a tufted appendix and a single diverticulum. Of 
two Abyssinian slugs described by Quick (1954), A.gughensis lacks caecum, appendix 
and diverticula but has a long, narrow, backwardly-directed sarcobelum ; while 
A. scotti lacks caecum, appendix and sarcobelum, but has two diverticula. In view 
of the intergradation and overlapping between species of which the anatomy is 
known, and of the number of species (especially in south-eastern Europe) of which 
it is not known, the use of sectional names within this genus does not seem useful. 



Agriolimax reticulatus (Miiller, 1774) 

Limax reticulatus Miiller, 1774 : 10. Denmark. 
Limax canariensis d'Orbigny, 1839 : 47. 
Krynickillus minutus Kalenickzenko, 1851 : 224. 

Notes 

The name agrestis has been used for this species by most British authors (see 
p. 170). 

First British record 
By Lister (1674 : 99) . 

Diagnostic features 

Distinguished from A. laevis and A. caruanae by larger size, opaque appearance 
with very variable coloration, white sticky mucus and less active movements ; 
and internally by the presence of a rectal caecum. Differs from A . agrestis in the trifid 
crenulate penial appendix, and the posterior situation of the ovotestis. 

External appearance (PI. 2, fig. 15) 

Length about 3-5 cm. The tail is obliquely truncated, the back keeled for a short 
distance at the hind end, and the sole tripartite. The respiratory orifice is at about 
two-thirds of the distance from front to back of the mantle, and the reproductive 
orifice close behind and below the right upper tentacle. The concentric striae of the 
mantle disappear in alcohol. The mucus is whitish on account of calcareous granules. 



BRITISH SLUGS 165 

The colour is variable, with the body usually marked with brown or grey. The 
respiratory orifice has a conspicuous pale border. 

External variation 

The colour varies from pale cream to bluish-black, but is usually some shade 
of brownish-yellow, with darker grooves and scattered dark brown markings of 
varied shapes and sizes. Named varieties and sub varieties are very numerous. 
Many are practically synonymous, while some of those listed by Taylor (1902-07) 
refer to other species. Individuals often become darker as winter approaches. 
The following varieties may be recognized : var. cineracea Moquin-Tandon (1855), 
entirely ash coloured ; var. violacea Gassies (1849), n ^ ac or slate-coloured ; var. 
rufescens Dumont & Mortillet (1856), rufous ; var. reticulata Muller (1774), grooves 
dark, giving a reticulate appearance ; var. nigra Morelet (1845), black, with pigment 
sometimes invading the sole. Albino forms occur occasionally. 

Shell (Text-fig. 10 f) 

The right border is convex. 

Pallial organs (Text-fig. 10 a) 

When the mantle is reflected to the right, a lobe of the kidney is seen to cover the 
rectum, and the short retractor penis to arise from the diaphragm at the anterior 
border of the pericardium. 

Retractor muscles (Text-figs, iig and h) 

The cephalic retractor muscles arise as a single stem in the mid-line, a short 
distance behind the diaphragm. The main stem usually divides basally, to form the 
right ocular retractor, and a common stem for the left ocular and the buccal bulb 
retractors (Text-fig. iig), but variations occur (Text-fig. n h). The genital retractor 
arises from the diaphragm near the front of the kidney, and is inserted between 
the vas deferens and the constriction above the sarcobelum. 

Nervous system 

This is very similar to that of Milax. 

Jaw 

Pale golden, of the usual oxygnathous type. 

Radula (Text-figs. 11 L and o) 

As in all Agriolimax, the endocones of the lateral teeth are delicate, lying close to 
the mesocone and on a slightly more superficial plane. The basal plates of the central 
and lateral teeth are larger and relatively narrower than those of A. agrestis. 

Table I compares the radulae of the four British species of Agriolimax. 



i66 





Fig. io. Agriolimax. Dissection with mantle reflected to the right. A. A. reticu- 
latus — (x 2), ovotestis extends much further backwards than in A. agrestis. B. 

A. agrestis — (x 2), ovotestis does not extend much further back than the rectal 
caecum. G. A. caruanae — (x 2-7), ovotestis far forwards, exposed on left side. 

No rectal caecum. D. A. laevis — (x 2-7). 

A. caruanae. E. Pallial region and cephalic retractors (x 12). K. Mating 

pair with sarcobela protruded. 

Shell (x 4). F. A. reticulatus. G. A. agrestis. H. A. caruanae. 

I. A. laevis. 



BRITISH SLUGS 



167 



Table I . — Radular Formulae of the British Species of Agriolimax Morch 

Remarks 



Locality- 



Date 



Formula 



Central tooth 
(mm) 



Agriolimax reticulatus (Miiller) 



Capel-y-mn, Brecknock 


20 . vii . 46 


C. 14. 22x86 


0-043x0-022 


— 




20.vii.46 


C.17. 27x107 


? 


— 


Blackpill, Swansea 


. 1 7 . viii . 46 . 


C. 14. 22x88 


? 


Small but 
mature. 


Mumbles Rd., Swansea 


. 23. viii. 46 


C.14.21X100 


0-035x0-017 


— 


Porteynon, Glamorgan 


. ? .xi.31 


C.16.22 


0-049x0-024 


— 


Clyne Valley, Swansea 


7 . ix . 46 


C. 17. 26x100 


0-043 XO-022 


— 


Llanwrtyd Wells 


. 18.xi.38 


C.6.8 




Young, 
8 mm long. 


Burpham, Surrey 


29 . xii . 46 


C. 18. 26x122 


0-052 x 0-026 


— 




Agriolimax agrestis (Linnaeus) 






Wheatfen Broad 


. 1 7 . viii . 46 . 


C.16.23 


0-038 xo-oi9 


Radula 
3-2 x 1 "7 mm 




23. viii. 46 


C.16.23X105 


O • 040 x • 02 


— 




1 7 . viii . 46 


C.14.21X100 


0-034 xo-oi7 


— 




? .x.46 


C.15.24 


0-038x0-019 


— 




? .ix.46 


C.15.? 


0-035 x 0-017 


— 




? 


C.15.21 X IOO 


0-032 xo-oi6 


— 




Agrioli) 


nax laevis (Miiller) 






Singleton, Swansea . 


10.iii.48 


C.13. 27x92 


0-023 XO-OI2 


Euphallic. 




2 . viii . 48 


C.13. 25x95 


0-023 X 0-OI2 


Hemiphallic 




Agriolima 


x caruanae Pollonera 






Sketty, Swansea 


. ? .x.31 


C.14.50 


— 


— 




? .iv.32 


C.14.55 


— 


— 


Singleton, Swansea . 


• 3i-i-35 


do.? 


0-023 xo-oi4 


Young, 
12 mm. 


Porteynon, Glamorgan 


? . vii . 46 


C.14. 55x118 . 


0-029x0-017 


Radula 
3-1 x 1-5 mm 



Alimentary system (Text-fig. 10 a) 

The tabulated salivary glands lie on each side of the crop, which narrows as it 
enters the small stomach. The posterior loop of the intestine extends behind the 
stomach towards the apex of the visceral mass, which is formed by the right lobe 
of the liver. The rectum has a small caecum overlying the ovotestis. 

Reproductive system (Text-fig. n c) 

The brown tabulated ovotestis lies towards the rear of the visceral mass, and is 
partly exposed on the surface (Text-fig. 10 a). The hermaphrodite duct is short 
and only slightly sinuous, with a seminal vesicle at its slender termination where it 
enters the albumen gland. The latter is short, shaped like a mammalian liver, and 

ZOOL. 6, 3. 12 



1 68 



BRITISH SLUGS 







c * 2± 



Fig. ii. Agriolimax. Distal genital ducts (x 5-3). A. A. laevis — euphallic indi- 
vidual. B. A. laevis — aphallic individual. G. A. reticulatus. D. A. agrestis. 

E. A. laevis — apex of penial appendage, showing bifid tip and two pilasters. 

Cephalic retractor muscles. F. A . laevis. G & H. A . reticulatus. I & K. 

A. agrestis. 

Representative radular teeth. L. A. reticulatus — (x 330). M. A. agrestis 

— (x 33°) • N. A. laevis — (x 330). O. A. reticulatus — (x 500) from young 
animal 8 mm long. 



BRITISH SLUGS 169 

pale brown in colour. The common duct, with the lobulated prostatic ribbon, is 
thrown into zig-zag folds. The free oviduct is short and straight, and the atrium 
small. The spermatheca duct arises from the atrium and expands to a narrow oval 
sac. The short vas deferens enters the penis beyond its apex, beneath the genital 
retractor muscle. The penis is narrow basally, expands to a segment containing 
the triangular sarcobelum, and narrows a little before the apical segment. The latter 
terminates in a large penial appendage which is trifid, each branch having a crenulated 
outline. The details of the branching vary to some extent. The branches, which 
are tubular, are everted and turned completely inside out during mating, as in 
A. agrestis and A. caruanae (see Text-figs. 12 d and e). 

Mating 

This can occur throughout the year in Britain, even in the winter months, taking 
place on the surface usually after dark. The pair follow one another for from half to 
one hour in a narrowing circle, with copious exudation of mucus. The sarcobelum 
is protruded and plays over the partner's body. Then the animals remain still, 
neck to neck with bodies curved, and each atrium and penis is everted, bringing 
the oviduct and spermatheca duct to the surface. Then the base of the penial 
appendix appears and is explosively turned inside out, the branches appearing as 
long thin hyaline threads. A mass of sperms enclosed in a ball of mucus is simul- 
taneously transferred from one partner to the other's everted mass, to which it 
adheres. The animals separate at once, and in a few seconds the genital complexes 
are invaginated and withdrawn, carrying the sperm packets with them. 

Development 

The eggs, laid about ten days after mating, are about 3 X 2-5 mm, translucent 
and speckled with calcareous dots which become absorbed during development. 
They hatch in two or three weeks, the young being 4 mm long, pale grey and trans- 
lucent, and like all the British species have no bands at any stage. The young grow 
rapidly, and in warm weather may mature in three months. During late embryonic 
development the caudal and cephalic vesicles can be seen pulsating through the 
translucent shell, when the eggs are viewed under water (see Text-fig. 18 g). In 
Britain several broods are produced by a pair, and 700 eggs may be produced in 
a season. Luther (1915) states that self-fertilization occurs in A. reticulatus isolated 
from birth. 

Ecology 

A. reticulatus is perhaps the commonest British slug. It prefers drier sites than 
A . agrestis and occurs in grass fields, root crops, hedges, gardens and rubbish heaps, 
and under stones and logs in woods and coppices. It is often a serious pest in gardens 
and allotments, devouring seedlings and green crops. 

Distribution (Map 15) 

Ubiquitous in the British Isles. Probably indigenous to most of Europe, the 



170 



BRITISH SLUGS 



Mediterranean region and the Atlantic isles, and introduced by commerce into 
most parts of the world. 

Fossil record 

Pleistocene to Recent. 

Material examined 

Living specimens from many places in England and Wales. Spirit material 
from Britain, Europe, Atlantic isles, Reunion, America and New Zealand. 



Agriolimax agrestis (Linnaeus, 1758) 

Limax agrestis Linnaeus, 1758 : 652. Sweden. 
Limax bilobatus Ferussac, 18 19 : 74. 
Limax pallidus Schrenk, 1848 : 143. 

Notes 

The commonest British slug, A. reticulatus (Miiller), was formerly regarded 
by British authors as a variety of A. agrestis, though Luther (1915) had shown 
them to be distinct. It is uncertain which of many synonyms in this group of slugs 
belong to A. agrestis and which to A. reticulatus. 

First British record 

By Ellis (1941). The slug was found at Wheatfen Broad, Norfolk, and determined 
by H. Watson. 

Diagnostic features 

Agrees externally with A. reticulatus, except that it is smaller, smoother and more 
slender and seldom has more than a few small dark flecks. Internally, the ovotestis 
lies further forward, and the penial appendix is short, smooth and unbranched. 

External appearance 

Like A. reticulatus, but rather slimmer and smoother, and less pigmented. The 
body is pale greyish-yellow, paler on the sides, sometimes with whitish calcareous- 
looking spots. 

External variation 

Occasionally, a few dark flecks are present and the grooves are slightly pigmented. 

Shell (Text-fig. 10 g) 

About 3 mm long, usually with the right border slightly concave. 



BRITISH SLUGS 



171 



Pallial organs (Text-fig. 10 b), retractor muscles (Text-figs. 11 1 and k), nervous 
system, jaw 

As in A. reticulatus. 

Radula (Text-fig. 11 m) 

The teeth are a little smaller than in A. reticulatus, and the basal plate wider 
(Table I, p. 167). 

Alimentary system (Text-fig. 10 b) 
As in A. reticulatus. 

Reproductive system (Text-fig. n d) 

The ovotestis lies further forward than in A. reticulatus, near the centre of the 
visceral mass, and is only partly exposed on the right side beneath the rectal caecum 
(Text-fig. 10 b) . The remainder of the reproductive system is like that of A . reticulatus, 
except for the apex of the penis, which terminates in a single small curved digit if orm 
appendix. 

Mating 

As in A. reticulatus. Mating occurs in Britain in the autumn, and perhaps at 
other times. Luther (1915) states that this species is incapable of self-fertilization, 
unlike A. reticulatus. 

Development 

The eggs are laid in clusters of from ten to twenty or more. They measure from 
2-2 X 2-o to 2-5 X 2-25 mm, and are more translucent and more finely dotted 
with calcareous particles than those of A. reticulatus. The newly hatched young 
are 3-5 mm long, translucent and whitish, and have no bands at any stage. 

In Britain, the eggs are laid in the autumn and hatch in three or four weeks. 
The young mature in the following summer and autumn, and die in the late autumn 
after mating. In Finland, the life cycle is different (Luther, 1915), in relation to 
the more severe climate. The eggs are laid, and the parents die, in late August or 
September, while the eggs do not hatch until the following June. Thus embryonic 
life lasts for more than eight months in Finland and less than a month in England, 
while post-embryonic life lasts about three and twelve months respectively. 

Ecology 

In Britain, apparently confined to marshy carr (see Ellis, 1941) in East Anglia. 

Distribution (Map 16) 

In Britain, certainly recorded only from marshes by the Norfolk Broads : a single 
record of an immature specimen from North Uist (Ellis, 195 1 : 196) has not been 
confirmed. Probably widespread in Europe, northwards to Finmark and Iceland, 



172 BRITISH SLUGS 

though the detailed distribution is uncertain because of confusion between this 
species and A. reticulatus. For example, Germain (1930) describes and figures 
A. reticulatus under the name agrestis. 

Material examined 

Living specimens from Wheatfen Broad, Sutton Broad and Alderfen Broad, all 
in east Norfolk. 

Agriolimax laevis (Muller, 1774) 

Limax laevis Muller, 1774 : 1. Frederiksdal, Denmark. 
Limax brunneus Draparnaud, 1801 : 104. 
Limax campestris Binney, 1844 : 52. 
Limax hyperboreus Westerlund, 1876 : 97. 
Agriolimax bevenoti Collinge, 1897c : 295. 

Notes 

Muller described his slug as " totum nigrum", except for the median area of 
the sole, and compared its general appearance with that of the black land planarian 
Rhynchodemus terrestris. However, topo types from Frederiksdal agree with the 
common slug, known as A. laevis in Britain, Europe and North America, in being 
some shade of brown. The discrepancy in colour is not highly significant, since the 
present species becomes very dark when somewhat dry and contracted, and since 
in America it varies in colour from amber to black (Pilsbry, 1948 : 540, 548 ; Altena, 
1958 : 30). 

Simroth (1885 : 222, pi. 9) described and figured, as forms of A. laevis from 
Germany, what are evidently representatives of two distinct species : those corre- 
sponding to the euphallic and aphallic forms of the widespread slug (his figs. 21 G 
and 22 h), and a larger and darker species with a hammer-headed penis (his figs. 
17 c and 18 d). The latter is found principally in Germany and Poland, though it 
appears to have been introduced to Gotland Island, Sweden (Lohmander, in litt.). 
It has never been recorded from Denmark and Lohmander, despite his considerable 
experience of Danish slugs, has never found it there. Taylor's figures (1902-07, 
figs. 131 and 132), purporting to show the genitalia of British A. laevis, are copied 
from Simroth and show this second species. 

It seems that the name laevis can properly be retained for the species to which 
it has long been applied. There is no reason to transfer it to the species with the 
hammer-headed penis, the correct name for which is uncertain. 

First British record 

By Johnston (1838 : 154), as Limax brunneus. 

Diagnostic features 

Small size, colour light or dark brown, smooth, active. Much smaller than 
A. caruanae. Head and neck extend further from the mantle than in A. reticulatus. 
Internally distinguished from the other species by its more sinuous penis, with a 



BRITISH SLUGS 173 

tubular caecum or appendix, and the frequent occurrence of aphallic specimens ; 
and from A. reticulatus and A. agrestis by the absence of a rectal caecum. 

External appearance (PL 2, fig. 19) 

Length 1-7-2-25 cm. Light or dark brown, translucent, sometimes greenish, more 
or less flecked with darker spots. The respiratory orifice is less conspicuously pale- 
bordered than in the other species. The head and neck are thrust further forward 
than in A. reticulatus and A. agrestis, so that the mantle is more centrally placed, 
and the mantle has fewer concentric ridges. The mucus is clear, not milky. The 
slug is very active, and crawls rapidly. 

External variation 

The ground colour varies from pale to dark brown, with more or less darker 
flecking on body and mantle. Pale and dark specimens occur together at all seasons. 
Named varieties are var. lacustris Bonelli (in Lessona & Pollonera, 1882) (= var. 
maculata Cockerell, 1886&), irregularly spotted with dark brown ; var. grisea Taylor 
(1904), grey or greenish-grey. 

There is also marked genital polymorphism, with euphallic, hemiphallic and 
aphallic individuals. 



Shell (Text-fig. 10 1) 

Relatively short and broad. 

Pallial organs (Text-fig. 10 d) 
As in A . reticulatus. 



Retractor muscles (Text-fig. iif) 

The cephallic retractor muscles arise by a long common stem, and so are less 
deeply divided than in the other species. The retractor penis is occasionally absent. 

Nervous system 

As in A . reticulatus. 

Jaw 

Pale golden, 1 mm across. 

Radula (Text-fig. iin) 

Rather fewer lateral teeth, and small ectocones on more of the external marginals, 
than in A. reticulatus and A. agrestis. Formula C.13.26. 

Alimentary system 

The rectum lacks a caecal diverticulum, as in A . caruanae. The visceral cavity is 
often somewhat pigmented, though less so than in A . caruanae. 



174 BRITISH SLUGS 

Reproductive system (Text-figs, n A and b) 

The dark ovotestis is less exposed than in the other species. The hermaphrodite 
duct is short and nearly straight, with the usual seminal vesicle at the short brown 
albumen gland. The female portion of the common duct is leaden grey, and the 
prostate yellow. The spermatheca is normally oval as in the other species, but 
sometimes in young examples it is globular. The penis in a euphallic individual 
shows four regions : a distal, narrower segment ; a wider one enclosing the triangular 
sarcobelum ; a twisted and contorted segment entered distally by the vas deferens, 
and giving attachment to the retractor penis ; and a short narrower segment 
(diverticulum or appendix), more or less bifid at the tip. The appendix contains 
two thickened pilasters, which fade out on reaching the sarcobelar segment. In 
the aphallic form (Text-fig. n b) prostate and spermatozoa are present, but the vas 
deferens fades out or ends in a little vesicle, before reaching a minute knob represent- 
ing the penis. Apparently the penis is always the last part to be fully developed ; 
but in young examples that will become euphallic the vas deferens does reach 
the small penis and a retractor muscle is often present. Stages can be found between 
this and the adult euphallic form. Babor (1894) records observations which he thought 
indicated that A. laevis was normally protogynous — young adults being always 
female and middle-aged adults hermaphrodite, with some old animals attaining a 
purely male phase, with atrophied albumen gland and spermatheca and hyper- 
trophied penis. He states that in the young stages sperms are very few, and there is 
no sperm duct or penis. However, dissection of many British examples, taken at 
various sites and different seasons, shows that here sperms, prostate and vas deferens 
are always present ; though in animals that will eventually be aphallic the vas 
deferens ends blindly. No example has been seen in which the albumen gland and 
spermatheca have atrophied. As these forms often occur together, in varying pro- 
portions in the same site at different seasons, it seems more likely that this is a case 
of genital dimorphism, such as occurs in Zonitoides. To establish this with certainty 
it is desirable that samples should be taken from the same site each month for a 
year, but an opportunity to do this has not occurred. Pilsbry (1948 : 539-552) 
gives numerous figures of similar dimorphism in the American forms of A. laevis. 

Mating 

Gerhardt (1939) says that following and circling around is omitted in A. laevis, 
and that the partners remain in contact before the sperms are transferred for a 
much longer period (up to an hour) than in the other species. 

Development 

The eggs are laid singly or in small clusters, probably at all seasons of the year, 
having been found in January, April and October. They are relatively large, from 
i-8 x 1-5 to 2 X 1*3 mm, translucent, resilient, finely punctate, with polygonal 
calcareous granules. They hatch in four or five weeks. The young are 4 mm long, 
white and translucent, with the opaque white liver lobes showing through the 
integument. The head and tentacles are light pinkish-brown with a violet tinge. 



BRITISH SLUGS 175 

After a few days, pigmentation begins at the tail and head, and gradually invades the 
rest of the body. 

Ecology 

A. laevis inhabits damper sites than our other slugs and is common in marshes 
and on the borders of ponds and rivers, but is occasionally found in drier sites, 
in fields and on the borders of woods. It has been seen feeding on mealy bugs infesting 
Streptocarpus in greenhouses. 

Distribution (Map 17) 

Probably ubiquitous in the British Isles. Its continental distribution is somewhat 
uncertain, because of the existence of a somewhat similar slug distinguished by a 
hammer-headed penis (p. 172). It is common in marshy places throughout the 
Holarctic, extending southwards in Europe to northern Italy. Most of this range 
is probably occupied by the true A . laevis, showing only slight infra-specific variation. 
A. laevis has been widely introduced into tropical regions and oceanic islands, where 
it has been recorded under many different names. For example, specimens from 
Hawaii in the B.M. (N.H.) labelled as A. bevenoti Collinge (Kauai) appear to belong 
to the aphallic form of this species, and others labelled as A . perkinsi Collinge (Lanai) 
and A. globosus Collinge (Mauna Loa) to the euphallic form. 

Fossil record 

Pleistocene to Recent. 

Material examined 

Living specimens from many places in England and Wales. Spirit specimens 
from the type locality (Frederiksdal, Denmark), kindly collected and sent to me by 
Dr. Mandahl-Barth, and from Britain, South America, New Zealand and Hawaii. 



Agriolimax caruanae Pollonera, 1891 
Agriolimax caruanae Pollonera, 1891 : 3. Porta Reale, Malta. 

Notes 

Pollonera described the keel of his species as blackish, and figured a more globular 
spermatheca than is found in adult British specimens. However, in Britain the keel 
and head are the first parts to pigment, so that in young animals they are darker 
than the rest of the body. The spermatheca is sometimes globular, in young animals 
at any rate in Britain, and in adults of some American forms. Pilsbry (1948 : 558) 
figures considerable variation between American specimens, which he nevertheless 
recognizes as belonging to A . caruanae. The British slug can be at most a geographical 
race of this species, of which Quick (1949 : 28) gives a short account. 



176 



BRITISH SLUGS 




^J&h^ 



Fig. 12. 



BRITISH SLUGS 177 

First British record 

Specimens, found by the late Charles Oldham in Cornwall about 1930, were 
identified with this species (as perhaps a distinct subspecies or variety) by H. Watson, 
and exhibited at meetings of the Conchological Society of Great Britain and Ireland 
and the Malacological Society of London. 

Diagnostic features 

Distinguished in life from the externally similar A. laevis by larger size and even 
greater activity. Internally distinguished by having two penial diverticula and a 
tuft of appendages, a darkly-pigmented visceral cavity, and a greater number of 
marginal radular teeth. 

External appearance (PL 2, fig. 17) 

The length when extended is 2-5-3 cm, or a little more. As in A. laevis, the head 
and neck extend far beyond the mantle. The slug is dark translucent chestnut 
brown, greyish-brown or grey, with the mantle lighter over the pallial region, and 
the body and mantle more or less flecked with dark brown. The respiratory orifice 
is pale rimmed. The sole is grey rather than brown, because the dark visceral pig- 
mentation shows through the tissues. The mucus is thin and colourless. 

External variation 

The colour varies from a clear chestnut-brown to a greyer brown, and the small 
darker flecks on body and mantle are more or less marked, but there are no named 
varieties. 

Shell (Text-fig. ioh) 

4-5 x 2 mm, or a little larger. In undistorted specimens, the right border is 
slightly concave. 



Pallial organs (Text-fig. 10 e) 
As in A . reticulatus. 



Fig. 12. Agriolimax caruanae. A. Reproductive organs (x 4). B. Penis 

( X 14) opened to show sarcobelum, two diverticula, entries of vas deferens and crenu- 
late branched appendix, penial retractor. C. Penis (x 8). D. Sarcobelum 

protruded, atrium everted, appendix beginning to be everted. E. Three branches 
of appendix fully everted, sarcobelum beginning to shrink. F. Distal genital 
ducts ( x 30) of young individual 1 1 cm long, showing oviducts, spermatheca, sarco- 
belum in penis, rudiments of diverticula and appendix. G & H. Cephalic retrac- 
tors. The buccal retractor is sometimes completely divided. I. Ganglia and 
commissures of nerve ring. K. Jaw (x 20). L. Representative radular 
teeth (x 500). M. Jaw ( x 100) of individual one day old. N. Jaw ( x 100) 
of young individual 6 mm long. 



178 BRITISH SLUGS 

Retractor muscles (Text-figs. 12 G and h) 

The cephalic retractors arise by a common stem, and the buccal retractor is 
divided either completely or for half its length. 

Nervous system (Text-fig. 12 1) 
As in A . reticulatus. 

Jaw (Text-fig. 12 k) 

Of the usual oxygnathous type, 1-3 mm across. In the newly-hatched animal, 
a median indentation (like that seen in Milax, Limax and Vitrina) reveals a bilateral 
origin (Text-figs. 12 m and n). 

Radula (Text-fig. 12 l) 

Characterized by having rather fewer lateral teeth and more marginals than 
A. reticulatus and A. agrestis. Formula C.14.55. 

Alimentary system (Text-fig. 10 c) 

As in A. laevis, there is no rectal caecum. The walls of the visceral cavity are 
darkly pigmented. 

Reproductive system (Text-figs. 10 c and 12 a-f) 

The dark ovotestis lies rather far forwards, and is partly exposed on the left side 
of the visceral mass. The oviduct, atrium and spermatheca require no special 
comment. The basal segment of the penis is slender and the segment containing 
the sarcobelum capacious. The distal segment bears apically two curved hollow 
processes, of which the left is often the longer, and between these an appendix 
with four or five long slender crenulated branches, corresponding with the apical 
appendix of A . reticulatus. 

Mating (Text-fig. 10 k) 

Breeding occurs throughout the year, eggs having been seen in January, May, 
June, July, October, November and December. A mating pair was observed on 
25th June, 1950. At 6.10 p.m. the partners were following one another in a circle, 
with protruded sarcobela, licking one another's tails. At 6.40 they were lying 
quietly with their necks close together, with the atria everted and the sarcobela 
shrinking. A few seconds later, the four long tubular branches of the penial appendices 
were everted, lying across the partner's body like hyaline threads. At 6.42 the 
everted organs were rapidly withdrawn, with the sarcobela last, and the animals 
crawled apart. 

Individuals from Glamorgan and Denbigh, isolated from birth, have on several 
occasions been seen to lay a few self-fertilized eggs, which have developed to an 
advanced stage although they have not been seen to hatch. Gregg (1944) reports 
fertile eggs from virgin specimens in California. There this species (for which Gregg 



BRITISH SLUGS 179 

uses the name Deroceras panormitanus) apparently passes through a purely female 
phase when about half grown. Only about half the individuals survive this phase, 
to grow to full size and develop male organs. There is no sign of such a female 
phase in Britain, where slugs only 10 mm long, with their common ducts still slender 
and immature, always have penes in which the rudiments of all the adult features 
are easily recognizable. Two diverticula, a budding tufted appendix and a sarcobelum 
can be seen in a penis between 1-3 and i-6 mm long. Aphallic and hemiphallic 
individuals are unknown in Britain. In any case, aphallic individuals of A. laevis 
are not in a female phase, since they have spermatozoa and a prostate. Gregg's 
observations are explicable in two ways : he may possibly have been working with a 
mixed culture of A . caruanae and A . laevis, or the difference may be related to the 
more rapid development of A. caruanae in California than in Britain. 

Development 

The eggs are relatively small, from 1-5 x 1*5 to 175 X 1-4 mm : much smaller 
than those of A . reticulatus and A . agrestis, and not much larger than those of A . laevis. 
They are laid in clusters of up to fifty eggs, and in the summer hatch in about seven- 
teen days. The young are only about 3 mm long, very pale and translucent, with 
pale violet tentacles. Pigmentation begins at the head and tail, as in A. laevis. 

Behaviour 

A . caruanae crawls extraordinarily fast for a slug, and is irritable and pugnacious, 
snapping at its neighbours and lashing its tail. Even when well supplied with food, 
it shows cannibalistic tendencies. 

Ecology 

Usually confined to gardens, in flower and vegetable beds, lawns, rubbish heaps 
and greenhouses ; but occasionally found at a distance from houses, in hedges, 
fields and waste places (records from Devon and Glamorgan). 

Distribution (Map 18) 

Recorded from scattered localities in England (mainly in the west), Wales, Scotland 
and southern Ireland (Makings, 1959). The extra-limital distribution is very imper- 
fectly known. Repeated efforts to obtain topotypes from Malta have failed. The 
species probably occurs at Marseilles (Pilsbry, 1948 : 560), and has been introduced 
into California. 

Fossil record 

Holocene shells from Romney Marsh (Hayward, 19540 and b) are very suggestive 
of this species. Although slug shells are badly characterized, especially when fossil, 
there is a prima facie case for believing that these are indeed shells of A. caruanae. 
Otherwise, the species is known only from Recent deposits. 

Material examined 

Living or spirit specimens from St. Agnes, Scilly Isles ; Exeter, Devon ; Swansea 



i8o BRITISH SLUGS 

and Porteynon, Glamorgan ; Old Colwyn, Denbighshire ; St. Albans, Hertfordshire ; 
Bromborough, Cheshire ; Edinburgh and Aberdeen, Scotland ; Cork (grounds of the 
University, from Dr. Makings) and Newcastle, Co. Down (Dr. Stelfox's garden), 
Ireland ; Pyrenees Orient ales ; Golden Gate Park, San Francisco. 



Genus LIMAX Linnaeus, 1758 

Type species Limax maximus Linnaeus, 1758. 

Limacinae in which the right ocular retractor passes forwards between the penis 
and the vagina. The intestine has two forwardly-directed loops, and the left lobe 
of the liver forms the apex of the visceral mass. There is no penial stimulator. 
The nucleus of the concentric mantle ridges lies in the mid-dorsal line. The dorsal 
keel slopes evenly to the end of the tail, which is therefore pointed and not truncated. 

Hesse (1926 : 8-16, 76-92) divides Limax into several subgenera, and the nominate 
subgenus into sections. Since no regular trends in several characters are discernible, 
each species being specialized in some respect or another (e.g. the long penis and 
peculiar mating behaviour of L. maximus and L. cinereoniger) , the infrageneric 
arrangement depends largely on the relative systematic importance attached to 
the radula, penis, alimentary tract and other organs. For example, L. nyctelius 
may be associated with L. flavus in the section Limacus, on the basis of its rather 
long cylindrical penis and long rectal caecum ; but their radulae are very different, 
that of L. flavus being specialized in the suppression of ectocones and endocones, 
while that of L. nyctelius approaches the condition of Agriolimax. Subgeneric 
and sectional names are therefore not adopted here. 



Limax tenellus Miiller, 1774 

Limax tenellus Miiller, 1774 : 11. Denmark. 

Limax tenellus', Nilsson, 1822 : 11. 

Limax serotinus Schrank, 1848 : 144. 

Limax cereus Held, 1849 : 15. 

Limax fulvus Normand, 1852 : 7. 

Agriolimax tenellus; Lessona & Pollonera, 1882 : 45, pi. 1. fig. 7. 

Notes 

Although Nilsson is often quoted as the author of this name, it was first applied 
by Miiller. Ignoring an obvious mistake in measurement, Muller's description seems 
adequate to identify the present species : " Limax virescens, capite tentaculisque 
nigris. Dan. Sp^ed-Sneglen. long. 10 unc. Totus albidus, Clypeus in luteum, 
abdomen in virescentem colorem aliquantum vergit ; ille margine postico, hoc 
apice supra nigricat. In Fossulis Nemorum foliis aridis repletis ; primo vere." 

Altena (1958) confirms that L. fulvus Normand is indeed L. tenellus. 

First British record 
By Alder (1848 : 124). 



BRITISH SLUGS 181 

Diagnostic features 

Distinguished externally from other British species by small size, and pale yellow 
colour, with black head and tentacles ; and internally by a short penis with no 
appendix. 

External appearance (PL 2, fig. 16) 

The smallest British Limax, from 25 to 35 mm long when extended. Colour 
pale yellow, or sometimes reddish-brown, with blackish head and tentacles, and 
sometimes with a pair of more or less distinct body and mantle bands. The mucus 
is yellow. 

External variation 

Var. cerea Held (1849), uniform yellow, or with mere traces of bands ; var. fulva 
Normand (1852), reddish-brown, suffused dorsally with black ; var. cincta Heyne- 
mann (1861), with distinct dark mantle bands, and sometimes body bands. 

Shell 

Thin and delicate, measuring about 3-5 x 2 mm. 

Pallial organs (Text-fig. 13 d) 

The somewhat angularly crescentic kidney partly embraces the heart and peri- 
cardium, just behind the origin of the penial retractor. 

Retractor muscles (Text-fig. 13 e) 

The cephalic retractor arises by two roots in the mid-dorsal line just behind the 
kidney, and divides about half-way along into right and left tentacular retractors. 
The right tentacular retractor passes forwards between the penis and oviduct, 
and not on the left of both as it does in Milax and Agriolimax. The pharyngeal 
retractor arises half-way along the left tentacular retractor, and divides into right 
and left branches passing to the buccal bulb between the cerebral and visceral 
ganglia. The penial retractor arises from the diaphragm on the left side, in front 
of the pericardium. 

Nervous system (Text-fig. 13 h) 

The olfactory and optic lobe of the cerebral ganglion is large, the cerebro-pleural 
and cerebro-pedal commissures short, and the visceral ganglia closely approximated, 
the right visceral and abdominal ganglia being fused together. 

Jaw (Text-fig. 13 f) 

Of the usual oxygnathous type, 1-5 mm across. 

Radula (Text-fig. 13 g) 

The central tooth, 0-038 x 0-018 mm, with small but distinct ectocones, is 
flanked on each side by about twenty lateral teeth with both ectocones and endocones, 



1 82 



BRITISH SLUGS 



followed by about forty-five marginal teeth. The teeth between the eighteenth 
and twenty-second are transitional in character, the endocone diminishing, and the 
ectocone approaching the mesocone. In the marginals, the mesocone becomes 
relatively longer and more curved, and the ectocone becomes prominent, arising 
from the side of the mesocone. Ectocones occur in some of the outer marginals 
of other Limax species, but they are not the conspicuous feature throughout that 
they are in L. tenellus. Formula C.20.45 x no. 




<3=\? 



G I 




10 iq 25 35 55 



Fig. 13. Limax tenellus. A. Reproductive organs (x 4). B. Penis (X 6-7) 

opened. G. Alimentary system, showing left lobe of liver forming apex of visceral 

mass, ovotestis at stomach level, between left and right lobes, intestine, cephalic 
retractor, oesophagus, salivary glands, buccal bulb, portion of albumen gland below 
left liver lobe. D. Kidney, heart and pericardium, origin of penial retractor in 

front, second forwardly-directed loop of intestine passing around cephalic retractor, 
rectum. E. Cephalic retractor muscles. F. Jaw ( x 14). G. Represent- 

ative radular teeth (x 330). H. Nerve ganglia (X 8). 



BRITISH SLUGS 183 

Alimentary system (Text-fig. 13 c) 

The left lobe of the liver forms the apex of the visceral mass. The apex of the 
stomach forms the most posterior part of the alimentary tract (which in Limax 
has almost lost the spiral torsion seen in Avion and in Milax), and the intestine has 
two forwardly directed loops. The first of these is held in place by the aorta as usual, 
and the second, which extends further forwards, is crossed by the cephalic retractor. 
The intestine runs backwards from this point and turns sharply to run forwards 
as the rectum, between the cephalic retractor and the floor of the pulmonary chamber, 
to the posterior lip of the respiratory orifice. 

Reproductive system (Text-fig. 13 a) 

The darkly pigmented ovotestis lies behind the stomach, but a little of it may be 
visible from the dorsal aspect, between the right and left lobes of the liver. The 
dilated central part of the hermaphrodite duct is pigmented and sinuous, narrowing 
at the albumen gland, where it has a distinct unpigmented vesicula seminalis. 
The prostate does not diverge anteriorly from the oviduct. The short vas deferens, 
which is not bound down by the right ocular retractor, becomes sacculate and pig- 
mented at the entrance to the penis, apically above the insertion of the penial 
retractor. Internally the penis (Text-fig. 13 b) shows three or four delicate short 
transverse folds apically, and further forwards three main massive longitudinal 
folds, the exact appearance varying according to where the penis is opened. These 
folds are visible during copulation (see below). The spermatheca is small and club- 
shaped, with a short duct entering the atrium. 

Mating 

According to Gerhardt (1933 : 445), one animal follows the other rapidly, head 
to tail. The leader soon bends to the right, so that after ten minutes the circle 
is completed. The penes are suddenly everted as massive bluish-white bodies, 
each with a thickened oblique fold which Gerhardt calls the comb. The penes are 
closely apposed, but not entwined, and the animals slowly revolve clockwise for 
one and a half hours. L. tenellus differs from L. maximus and L. cinereoniger in 
not entwining the body nor hanging from a horizontal surface. 

Development 

The eggs vary from 3-6 x 3-1 to 3-3 x 3-0 mm, and are pale amber, soft, trans- 
lucent and slightly adherent. In Britain, they are laid from November to March 
in clusters of about fifteen, and take from sixty to 120 days to hatch. The young 
are 6 mm long, with a slight dorsal keel posteriorly. They are white and translucent, 
without body and mantle bands and with a more opaque kidney region. The tentacles 
are pale violet, with the dark brown retractor muscle showing through. 

Ecology 

Restricted to woodlands, usually of considerable extent, and one of the few 
slugs found in pine woods. Absent from most of the many woods and plantations 

ZOOL. 6, 3. 13 



1 84 BRITISH SLUGS 

established during the seventeeth century, which suggests that its powers of dispersal 
are poor. Indifferent to the presence or absence of lime. 

Distribution (Map 19) 

Of very local occurrence, though wide range, and therefore seldom encountered. 
In Britain, recorded from many vice-counties. In Europe, found from France 
northwards to Jemtland, and eastwards to the Caucasus. 

Material examined 

Living specimens from Netley Heath, Shere, Surrey ; and Chepstow, Monmouth- 
shire. Spirit specimens from Britain and Switzerland. 



' Limax flavus Linnaeus, 1758 

Limax flavus Linnaeus, 1758 : 652. Sweden. 
Limax variegatus Draparnaud, 1801 : 103. 
Limax megalodontes Quoy & Gaimard, 1824 : 426. 

First British record 

By Lister (1685-97, pi. 101b). 

Diagnostic features 

Tentacles steely blue, body and mantle with yellowish spots and no lateral bands. 
Long rectal caecum and long cylindrical penis — a diagnostic combination for British 
Limax. 

External appearance (PI. 2, fig. 22) 

A rather large slug, from 7 to 10 cm long when extended. Colour from yellowish 
to greenish, more or less suffused with dusky pigment, and mottled on the mantle 
with yellow patches, on the body with yellow spots formed by groups of up to seven 
tubercles. There are no lateral bands. The tentacles are a very characteristic cold 
steely blue, the sole pale yellow, the respiratory orifice pale-rimmed, and the mucus 
yellow. 

External variation 

Var. rufescens Moquin-Tandon (1855), reddish-brown ; var. virescens Ferussac 
(1819), uniformly greenish ; var. antiquorum Sowerby (1834 ?-)» P a * e ochraceous 
marbled with grey ; var. breckworthiana Lehmann (1864), uniformly suffused with 
dark pigment. 

Shell 

Rather large, 9x6 mm, thin and convex, with the nucleus near the posterior 
margin and a little to the left of the mid-line. 



BRITISH SLUGS 



185 



Pallial organs 
As in L. tenellus. 



Retractor muscles (Text-fig. 14 d) 

The cephalic retractors arise from two roots, which do not unite for a considerable 
distance. The buccal retractor arises near the origin of the left ocular retractor. 
The penial retractor arises from the diaphragm, in the region of the kidney. 

A c B 




Fig. 14. Limax flavus. A. Alimentary system ( x 2) showing second forwardly- 
directed loop of intestine passing around cephalic retractor, long rectal diverticulum 
passing backwards. B. Reproductive organs (x 2). G. Penis (x 6) opened 

to show internal folds. D. Cephalic retractor muscles. E. Jaw (x 14). 

F. Representative radular teeth ( x 330). 



186 BRITISH SLUGS 

Nervous system 
As in L. tenellus. 

Jaw (Text-fig. 14 e) 

2-25 mm across, dark brown in colour. Marked transversely with fine striae, 
which are crossed by still finer ones following the contour of the jaw. 

Radula (Text-fig. 14 f) 

Measures 6-5 X 2-6 mm, with about 150 rows of teeth. The central tooth (measur- 
ing 0-05 X 0-038 mm) has a strong mesocone, but its ectocones are obsolete or 
absent. The lateral teeth are also devoid of ectocones and endocones. The aculeate 
marginals, some of the outermost of which have small ectocones, are connected 
to the laterals by several transitional teeth. Formula C. 16 4-4.50 X 150. 

Alimentary system (Text-fig. 14 a) 

The oesophagus dilates to a capacious crop, flanked by the salivary glands, and 
merging gradually into the stomach. The second forwardly directed loop of intestine 
is held in place by the cephalic retractor muscle. Instead of being prolonged back- 
wards as it is in L. tenellus, it soon turns forwards as the rectum ; but from the apex 
of the bend a long rectal diverticulum extends backwards, on the surface of the liver, 
to the end of the body. As in the other species of Limax, the left liver lobe forms 
the apex of the visceral mass. 

Reproductive system (Text-fig. 14 b) 

The unpigmented ovotestis lies between the lobes of the liver, well below the 
stomach apex. The hermaphrodite duct, at its entrance to the yellow linguiform 
albumen gland, has a rounded white seminal vesicle. The ribbon-like yellow prostate 
diverges from the oviduct anteriorly, and can easily be peeled off it for the greater 
part of its length. The vas deferens runs forward, and turns back beneath the right 
ocular retractor to the apex of the penis. The penis is a rather long cylindrical 
organ, strongly folded in a spiral. Internally it bears (besides three or four minor 
longitudinal folds) a long, low, crinkled, ribbon-like fold along its whole length, 
and another more prominent fold that expands distally to a crest or " comb ", 
lobulated or crenulated on the free margin (Text-fig. 14 c). The oviduct expands at 
its entrance to the atrium, where the spermatheca arises from it. 

Mating 

Mating occurs from late summer to February. According to Gerhardt (1933), 
after " following " for about eight minutes, the animals approximate the right 
sides, but do not curve into a circle. The penes are rapidly everted and entwine 
spirally, forming a mass of about one square centimetre in area. Sperms enclosed 
in a mucous mass are transferred between the penes, which are rapidly withdrawn 
carrying the partner's sperms with them. The whole process occupies only half 



BRITISH SLUGS 187 

a minute. Gerhardt does not state what follows, but presumably the sperms are 
transferred from the withdrawn penes to the spermathecae. 

Development 

The eggs are laid in clusters of up to twenty, and have been found in September 
and February. They are large, 6x4 mm, pale amber in colour, and differ from 
those of other British slugs in having a small rounded projection at each end, so 
that they resemble miniature lemons. They hatch in from three to six weeks at 
room temperatures. The newly-hatched young are from 10 to 13 mm long, pale 
greenish-yellow with the characteristic blue tentacles already evident. There are 
no bands at any stage, and the yellow spots of the adult appear later in development. 

Behaviour 

L. flavus is essentially nocturnal, and unless looked for after dark it may escape 
observation in urban surroundings, even where it is abundant. 

Ecology 

L. flavus is found in woods, under logs and bark, but is much commoner in gardens, 
in the crevices of walls, and in cellars and out-houses. It eats decaying vegetable 
matter, fungi and lichens in wild places, and mildew and garbage in gardens and 
out-houses. 

Distribution (Map 20) 

Probably occurs in every vice-county of the British Isles, and ranges from southern 
Norway to North Africa and eastwards to Syria. It has been introduced into South 
Africa, Australia and North and South America, and to islands in the Atlantic 
and Pacific. 

Material examined 

Living specimens from Swansea, Glamorgan ; and Reading, Berkshire. Spirit 
specimens from Britain, Corsica, Greece, North and South America, South Africa, 
Australia, and the New Hebrides and Cook Islands. 



Limax cinereoniger Wolf, 1803 

Limax cinereoniger Wolf, 1803 : 7. Germany. 
Limax antiquorum F6russac, 18 19 : 68 (in part). 
Limax maximus; Gray, 1840 : 113 (in part). 
Avion lineatus Dumont, 1850 : 64. 
Limax cinereus var. intermedia Breviere, 1881 : 314. 

First British record 
By Gray (1840). 



188 BRITISH SLUGS 

Diagnostic features 

Largest British slug, usually black with dorsal line and median zone of sole white, 
keel prominent and rather long. Distinguished from L. maximus by coloration, 
and by coarser tubercles, spotted tentacles, relatively small shell and jaw, and longer 
penis not tapering distally. 

External appearance (PL 2, fig. 25) 

This is our largest native slug, which grows to 10 or 20 cm long in this country, 
while some continental forms are even larger. The keel is prominent and long, 
and the tubercles relatively coarser than in L. maximus. The slug is usually black 
all over, except for the white mid-dorsal line and median area of the sole. The dark 
tentacles are spotted with small discrete black or dark brown dots — a character 
distinguishing the species from L. maximus, which has apparently not been recorded 
previously. 

External variation 

In Britain the ground colour, seen in the mid-line and the median area of the sole, 
is usually whitish, but in continental specimens the keel may be red or yellow. The 
following named varieties have been recorded from Britain : var. ornata Lessona 
(1880), body black with a series of white spots along each side ; var. maura Held 
(1836), entirely black ; var cinerea Moquin-Tandon (1855), entirely ash-coloured 
except for bluish-black mantle ; var. punctata Lessona (1880), keel and mid-line 
of dorsum yellow, two interrupted bands on each side. 

Shell 

Relatively smaller (9x5 mm), thinner and slightly more elongated than that 
of L. maximus. 

Pallial organs 
As in L. tenellus. 

Retractor muscles 

The slender cephalic retractor arises from the mid-line behind the diaphragm, 
and crosses the second forwardly-directed intestinal loop. The right ocular retractor 
passes (as is usual in Limax) between the male and female divisions of the distal 
genital ducts, over the vas deferens. The penial retractor arises broadly from immedi- 
ately to the left of the cephalic retractor, and narrows to its insertion at the apex 
of the penis. 

Nervous system 
As in L. tenellus. 



BRITISH SLUGS 



Jaw (Text-fig. 15 e) 

Usually about 3 mm across, but may reach 4 mm in very large specimens 
relatively smaller than in L. maximus. 




Fig. 15. Limax. A. L. cinereoniger — distal genital ducts ( x 1-3). B. L. maxi- 

mus — mating pair suspended from mucus thread (adapted from L. Adams). G. 
L. maximus — distal genital ducts (x 2). D. L. maximus — stages in eversion and 

entwining of penes (adapted from L. Adams). E. L. cinereoniger — jaw (x 6-7). 

F. L. maximus — jaw (x 6-7). G. L. cinereoniger — representative radular teeth 

( x 33°) ■ H. L. maximus — representative radular teeth (x 330). 



i go BRITISH SLUGS 

Radula (Text-fig. 15 g) 

There are about 150 rows of teeth. The central tooth has rather small but strong 
ectocones. It is flanked by about nineteen laterals, with ectocones and endocones, 
the latter descending on the mesocones in the outer teeth. There are about fifty 
marginals on each side, of which only the inner teeth retain the endocone. In the 
outer marginals, an ectocone appears on the concave side of the mesocone, and 
becomes bifid or trifid in a few of the outermost teeth. Formula C.19.50 x 150. 

Alimentary system 

Resembles L. tenellus, and differs from L. flavns, in having the loop of intestine 
which is retained by the cephalic retractor prolonged backwards almost to the 
visceral apex, and without a rectal caecum. The loop is longer than in L. maximus. 

Reproductive system (Text-fig. 15 a) 

The long narrow darkly pigmented ovotestis is visible on the surface between 
the left and right liver lobes. The hermaphrodite duct is pale, slender at first, 
then wider and folded, and narrows at the somewhat triangular albumen gland. 
If a seminal vesicle is present, it is small, deep in the albumen gland and difficult 
to find. The folded common duct narrows to a long free oviduct, dilated at its 
entry into the atrium, and the small club-shaped spermatheca arises one or two 
millimetres before its end. The prostate separates from the common duct and so 
is free at its anterior end, and the vas deferens passes under the right ocular retractor 
and back to the apex of the penis. The stout cylindrical contorted penis is of nearly 
uniform diameter throughout its length, and when straightened out is from 6-8 cm 
long. Internally it bears a prominent fold, double at its origin. In its upper part 
this expands into a prominent frill, the " comb ". The rest of the mucous membrane 
is circularly ridged. 

Mating 

Mating occurs at night, throughout the milder periods of the year, on vertical 
or overhanging surfaces. After about fifteen minutes of " following ", the animals 
hang down freely from a disk of mucus. Their bodies and the evaginated penes, 
which become as long as the bodies, are entwined. A mass of sperms enclosed in 
mucus descends each vas deferens inside the penis, and the sperm packets are 
exchanged between the "combs" which are now terminal. The penes are then 
withdrawn, carrying the sperms with them to be transferred later to the spermathecae. 
The whole process takes about twenty minutes. 

Auto-fecundation can occur in this species (Oldham, 1942a). 

Development 

The eggs are about 5 X 5*5 mm, soft, translucent and amber-coloured. On account 
of the large size of the eggs, and their transparency when under water, the caudal 
and cephalic vesicles of the embryos can easily be observed (Text-fig. 18 h). The 



BRITISH SLUGS 191 

eggs hatch in about a month. The newly hatched young are translucent white, 
but soon become opaque, then brown and finally black. There is sometimes a sug- 
gestion of banding, and the lateral areas of the foot are the last to pigment. 

Oldham (1942&) records that this slug becomes mature in two years, and that an 
individual in captivity lived for five years and 270 days. 

Ecology 

This is essentially a slug of wild places, being found beneath logs and leaves and 
in tree crevices of ancient woodlands. In Pembrokeshire at least, it also occurs on 
damp lichen-covered rocks on bare hillsides. It seems never to occur in gardens and 
cultivated land or recent plantations. Though in the wild it presumably feeds on 
lichens, fungi and decaying vegetable matter, in captivity it readily eats carrot, 
oatmeal etc. 

Distribution (Map 21) 

L. cinereoniger is recorded from most British vice-counties, except for East Anglia 
and parts of central England, southern Scotland and central Ireland. In Europe, 
it extends from arctic Norway to north-eastern Spain, Italy, Sardinia, Greece and 
the Crimea. 

Material examined 

Living specimens from Netley Heath, Surrey ; Cusop Dingle, Hereford ; Buckle- 
bury, Berkshire ; and Pont Neath Vaughan, Glamorgan. Spirit material from Britain, 
Denmark, Germany and Luxemburg. 



Limax maximus Linnaeus, 1758 

Limax maximus Linnaeus, 1758 : 652. Sweden. 

Limax cinereus Muller, 1774 : 5. 

Limax antiquorum Ferussac, 18 19 : 68 (in part). 

First British record 
By Lister (1674 : 99). 

Diagnostic features 

Large, differs from L. cinereoniger in coloration : grey or brown with two or three 
bands on each side of body, dark spotting or marbling on mantle, tentacles pale 
pinkish-brown without spots, sole uniformly pale ; and also in having the tubercles 
relatively smaller, the shell and jaw relatively larger and the penis shorter and 
tapering distally. 

External appearance (PL 2, fig. 24) 

Length from 10 to 20 cm. The tubercles are relatively smaller than in 
L. cinereoniger. The slug is usually some shade of grey, with a uniformly pale sole. 



192 BRITISH SLUGS 

The body bears two or three bands, often doubled or more or less interrupted, on 
each side. The mantle is irregularly spotted or marbled with dark pigment, but 
never banded. The tentacles are translucent pinkish-brown, without spots. 

External variation 

The ground colour may be grey, ochraceous or reddish, and the bands may be 
developed to an infinitely variable degree. Named varieties are : var. concolor 
Pini (1876), uniformly ash-coloured ; var. Candida Lessona & Pollonera (1882), 
white, translucent, eyes pale brown ; var. vinosa Baudon (1884), purple or vinous 
brown ; var. fasciata Razoumowsky (1789), ash-coloured with three bands on each 
side ; var. sylvatica Morelet (1845), ash-coloured with two bands, and an indistinct 
third lower band ; var. tetrazona Taylor (1902), with only two bands on each side ; 
var. cellaria Dezallier d'Argenville (1742), bands interrupted; var. aldrovandi 
Moquin-Tandon (1855), ash-coloured with pale spots ; var. tigris Taylor (1902, 
from Adams MS.), tawny yellow with black bands. 

Shell 

Measures about 11x7 mm, and is thus relatively larger than that of L. cinere- 
oniger. 

Pallial organs, retractor muscles, nervous system 
As in L. cinereoniger. 

Jaw (Text-fig. 15 f) 

Relatively larger (3-5 mm across) than that of L. cinereoniger. 

Radula (Text-fig. 15 h) 

The central tooth is larger and relatively narrower than in L. cinereoniger, and 
its ectocones are more delicate and sometimes almost obsolete. There are about 
nineteen lateral teeth on each side, all or most of which retain both ectocones and 
endocones. The marginals (about fifty on each side) lack endocones, and only some 
of the outermost teeth have small ectocones. Formula C.19.50. 

Alimentary system 

As in L. cinereoniger, except that the last posteriorly-directed loop of intestine 
is shorter, reaching only about half-way along the visceral mass. There is no rectal 
caecum. 

Reproductive system (Text-fig. 15 c) 

The ovotestis is of the same long flattened linguiform shape as in L. cinereoniger, 
but is pale or only slightly pigmented. The hermaphrodite duct, albumen gland, 
common duct, free oviduct and spermatheca are also similar. The distal end of 
the prostate diverges more or less from the oviduct, and above this the prostate is 



BRITISH SLUGS 193 

only weakly attached for a considerable distance. The penis differs from that of 
L. cinereoniger in being shorter, widest apically and tapering distally. Internally 
there is a fold, expanded proximally into a " comb ", as in L. cinereoniger. 

Mating (Text-figs. 15 B and d) 

This occurs at night from overhanging surfaces. Unlike L. cinereoniger, L. maximus 
hangs suspended in mid-air from a stout thread of mucus. The protrusion of the 
organs, sperm transfer, and withdrawal of the penes is as in L. cinereoniger, but the 
entwined penes are not so long. After withdrawal, the animals re-ascend the mucus 
thread. Adams (1898) gives further details. 

Development 

The eggs, 5-0 x 5*5 mm, resemble those of L. cinereoniger. They are laid in early 
spring and in the autumn. February eggs in an unheated room hatch in six and a 
half weeks. The young are pale grey, with the middle lateral bands visible from 
hatching — thus differing from young Lehmannia marginata, in which the upper 
band appears first and mantle bands are also present. The tentacles are pale pinkish- 
grey. After two or three weeks, the other body bands have appeared and begin 
breaking up, and the mantle becomes marbled. 

L. maximus lives for at least three or four years. 

Ecology 

L. maximus occurs in woods, hedgerows and waste places, where it is sometimes 
found together with L. cinereoniger ; but unlike the latter it occurs also in gardens, 
cellars and out-houses. It seems to avoid green leaves and to feed on fungi and 
decaying matter, though in captivity it readily eats oatmeal, Bemax etc. 

Distribution (Map 22) 

Apparently general in the British Isles, and found from Oslo and Petrograd 
to Asia Minor, Algeria and the Atlantic isles. Introduced into North America, South 
Africa, Australia, Tasmania and New Zealand. 

Material examined 

Living specimens from Swansea, Glamorgan ; Savernake Forest, Wiltshire ; 
Bucklebury Common, Berkshire. Spirit material from Britain, France, Greece, 
Azores, Madeira, Canary Isles and British Columbia. 



Genus LEHMANNIA Heynemann, 1861 

Type species Limax marginatus Muller, 1774. 

The penis is short, with a laterally-placed apical appendix (conical in L. marginata, 
short and cylindrical in L. poirieri, and long and cylindrical in L. melitensis Lessona 
& Pollonera). A long rectal caecum extends to the apex of the visceral mass. In 



194 BRITISH SLUGS 

L. poirieri and L. melitensis the radula is like that of Agriolimax ; but that of 
L. marginata is highly specialized. The body readily absorbs water and becomes 
swollen and translucent. 

Hesse (1926) adopts sectional names within Lehmannia, segregating L. marginata 
(Lehmannia s.s.) from L. poirieri and L. melitensis (Ambigolimax) because of the 
former's specialized radula, although their reproductive and alimentary systems 
are very similar. As for Limax, subdivision of the genus depends on the importance 
ascribed to different characters. 



Lehmannia marginata (Miiller, 1774) 

Limax marginatus Miiller, 1774 : 10. Denmark. 
Limax arborum Bouchard-Chan tereaux, 1837 : 164. 

First British record 

By the Reverend B. J. Clarke (in Thompson, 1840 : 204). 

Diagnostic features 

Gelatinous aspect in damp weather, mantle bands forming lyre-shaped figure, 
two body bands on each side ; conical penial appendix, rectal caecum, specialized 
radula. 

External appearance (PL 2, figs. 18 and 21) 

Usually about 7-5 cm long, or larger. Its great capacity for absorbing water 
makes this slug become semi-transparent and gelatinous in appearance during damp 
weather. When irritated, it exudes copious watery mucus. In colour it is character- 
istically grey with a pale sole, two darker body bands on each side (which may be 
more or less indistinct or interrupted), and a pair of mantle bands forming an 
elongated lyre-shaped figure. The keel is paler than the body, and the median area 
of the mantle often darker. 

External variation 

Var. glauca Clarke (1843), ground colour greenish-grey ; var. subrufa Le Compte 
(1871), ground colour yellow; var. rosea Broeck (1870), ground colour pinkish; 
var. bettonii Sordelli (1870), body bands fused together and interrupted; var. tigrina 
Wienland (1876), with rows of black spots on body and mantle ; var. rupicola 
Lessona & Pollonera (1882), darkly pigmented, with indistinct markings. 

Shell 

White and iridescent. Normally measures 4 X 275 mm, but varies considerably 
in size and thickness. 

Pallial organs 
As in Limax. 



BRITISH SLUGS 



195 



Retractor muscles (Text-fig. 16 c) 

The main stem of the cephalic retractors arises from the mid-dorsal line just 
behind the kidney, and divides about half-way into the right and left ocular retractors. 
The buccal retractor arises from the left ocular retractor, a short distance in front 
of the bifurcation. The strong penial retractor arises from the diaphragm between 
the pericardium and the posterior border of the kidney, to be inserted below the 
vas deferens. 




C I 





UO 50 51 10 




Fig. 16. Lehmannia marginata. 
( X 5-3) opened longitudinally, 
tract and saliva ry glands. E. 

(X 330). 



A. Reproductive organs (x 3-3). B. Penis 

G. Cephalic retractor muscles. D. Alimentary 

Jaw ( x 13). F. Representative radular teeth 



ig6 BRITISH SLUGS 

Nervous system 
As in Limax. 

Jaw (Text-fig. 7 e) 

1 -4 mm across, pale yellow in colour, and of the usual Limacid shape, not as narrow 
as figured by Taylor (1902-07, fig. 106). 

Radula (Text-fig. 16 f) 

Very characteristic, and easily distinguishable from the radulae of other British 
slugs. The central tooth has a strong, broad and short mesocone, and very small 
or obsolescent ectocones. It is flanked by ten or eleven laterals, with short wide 
mesocones and no ectocones, of which only the innermost one or two may have 
small endocones. The numerous marginals (up to seventy on each side) have charac- 
teristically strong, arched reflections bearing the short wide mesocones. From about 
the fortieth marginal outwards, the mesocone becomes bifid and then trifid, and 
minute denticulations corresponding to ectocones appear on the reflection. 

Alimentary system (Text-fig. 16 d) 

Resembles that of Limax flavus, with a rectal caecum extending backwards on 
the surface of the liver to the level of the stomach apex. The visceral cavity is 
darkly pigmented. 

Reproductive system (Text-fig. 16 a) 

The ovotestis is bulky and compact and superficially pigmented. The hermaphro- 
dite duct is pale, swollen and folded in its median portion, and bears a rounded 
seminal vesicle where it terminates at the short linguiform albumen gland. The 
oviduct and spermatheca resemble those of Limax. The prostate is extraordinarily 
massive and deeply pigmented. The vas deferens, bound down by the right ocular 
retractor muscle, is short and wide, and enters the penis at the side of its blunt 
apex. At the opposite side of the apex there is a short conical appendix. The penis 
opened longitudinally (Text-fig. 16 b) shows a prominent fold, expanded and free 
at the apex and almost smooth, but corresponding with the " comb " of Limax. 
There is a small conical fold at the entrance to the appendix. The walls of the 
proximal part of the penis are thick and rugose. 

Mating 

This species can mate on level, vertical or overhanging surfaces. 

Development 

In Britain, mating occurs during the winter months. December eggs in an unheated 
room hatch in February. The eggs are soft, very translucent and pale amber in 
colour. They usually measure 4-3 x 3-6 mm, but some are smaller. The young 
are 10 mm long, pale translucent grey with violet tentacles. They show well-marked 



BRITISH SLUGS i< J7 

lyriform mantle bands, and a distinct band high up on the body — in contrast to 
young Limax maximus, in which the middle band is the first to appear. 

Behaviour 

Like many (perhaps most) slugs, especially when young, L. marginata is able 
to suspend itself from a mucus thread secreted by the pedal gland. This thread 
passes back along the foot to the tip of the tail and adheres to a branch or other 
object, and supports the animal when it crawls off. Sometimes the slug can even 
turn around and re-ascend the thread. Kew (1902) gives numerous examples of 
this process, with details and some figures. 

Ecology 

Not uncommon in old and newer woodlands, especially in the west and north. 
In dry weather it retreats into crevices in the bark and between roots, while in 
damp weather it crawls up the trunks to a height of twelve feet or more. It is also 
found on damp rocks on exposed hillsides, and on old stone walls. 

Distribution (Map 23) 

Probably occurs in every vice-county of the British Isles, but is commoner in 
the west and north. In Europe it is found from Iceland and Lapland to Russia 
and Italy, and is recorded as introduced into Australia and New Zealand. In North 
America, it apparently occurs wild at Torbay, Newfoundland (confirmed by Altena, 
1950). However, introduced slugs from Verdugo Woodlands, Glendale, California, 
kindly sent by Dr. W. O. Gregg and recorded as this species, proved to be L. poirieri. 
Probably some of the records from southern Europe also refer to L. poirieri. 

Material examined 

Living specimens from the Gower Peninsula and the neighbourhood of Swansea, 
Glamorgan ; Brecknock ; Caernarvon ; and Torrington, Devon. Spirit material 
from Britain and Australia. 



GREENHOUSE ALIENS 

Lehmannia poirieri (Mabille, 1883) 

? Limax valentianus Ferussac, 1823, 2 : 96 e . 
Limax poirieri Mabille, 1883 : 52. Spain. 

Notes 

A full list of references is given by Altena (1950 : 9-17). 

First British record 

By Quick (1949 : 24), as L. valentianus. However, Dr. A. D. J. Meeuse had already 
found it in a greenhouse in Belfast Botanical Gardens, on 26th November, 1948. 



i 9 8 BRITISH SLUGS 

Diagnostic features 

Somewhat like L. marginata, but the ground colour is yellower and paler, and the 
body bands nearer to the mid-dorsal line. The radula is like that of Agriolimax, 
very different from the specialized radula of L. marginata. The penial appendix 
is longer and more cylindrical than that of L. marginata, and not pointed at the apex. 

External appearance (PL i, fig. 14) 

L. poirieri is about 6 cm long, with a yellowish-grey body and mantle, and slightly 
darker head. The general appearance is watery and translucent, like a pale L. 
marginata. The keel is short and inconspicuous. A pair of body bands, high up 
near the mid-dorsal line, are usually more or less interrupted and with scalloped 
outer edges. Sometimes faint traces of a second pair of bands occur lower down. 
A pair of mantle bands, becoming broader and darker behind, form a lyre-shaped 
figure, and the area enclosed is often darker in the centre and mottled with brown. 
The respiratory orifice is pale bordered. The tripartite sole is uniformly pale. 

External variation 

The mantle and body bands vary in width, completeness and the degree of crenu- 
lation of their outer borders, and there are sometimes faint traces of a second pair 
of body bands. 

Jaw 

Pale yellow, 2-25 mm across. 

Radula (Text-fig. 17 d) 

Like that of Agriolimax in having a distinctly tricuspid central tooth, and endo- 
cones on the lateral teeth — a feature which led Pollonera (1 887*2 : 2) to place the 
species erroneously in that genus. The tri-cuspid central tooth is flanked by fifty-five 
to sixty teeth on each side, of which about fifteen are typical laterals with endo- 
and ectocones. Most of the inner marginals are simply aculeate, but many of the 
outer ones are bifid, while in the extreme outer ones the main cusp becomes multi- 
denticulate. 

Alimentary system 

The intestine shows the two forwardly-directed loops and the long rectal caecum 
reaching to the apex of the visceral mass, as in L. marginata and Limax flavus. 

Reproductive system (Text-fig. 17 a) 

Similar to that of L. marginata, but with the penial appendix longer, and bluntly 
cylindrical rather than conical. Pollonera (18870) figures the appendix of valentianus 
as dilated and terminally vesicular, but this may result from misinterpretation of a 
somewhat folded appendix (Simroth, 1887). Altena (1950) found that Canadian 



BRITISH SLUGS 



99 



specimens had simple cylindrical appendices, like those of specimens from Swansea. 
Another difference from L. marginata is that the sac into which the spermatheca 
duct expands is somewhat sausage-shaped, rather than pyriform. 




Fig. 17. Greenhouse aliens. A. Lehmannia poirieri — genital organs (x 3-3). 
B. Limax nyctelius — genital organs (x 3-3). G. L. nyctelius — penis (x 6-7) 

opened longitudinally. D. L. poirieri — representative radular teeth (x 330). 

E. L. nyctelius — representative radular teeth (x 330). 

zool. 6, 3. 14 



200 BRITISH SLUGS 

Mating, development 

No observations on mating seem to have been recorded. Eggs laid in captivity 
at Swansea in December 1939 were translucent yellow and measured 2-25 X 1*5 
mm. They hatched in February. In March the young measured io-o mm long, 
and had intensely black body and mantle bands, and by 10th June they were full 
grown. 

Ecology 

Found always on the ground, never climbing trees and plants like L. marginata. 
This is true in Tenerife (Altena, 1950) and California (Gregg, in litt.), and in a green- 
house at Swansea, where the species was always found under pots. 

Distribution 

Spain, the Canary Isles (? introduced), and introduced into California and Britain 
(known from greenhouses in Reading, Swansea and Belfast) . 

Material examined 

Living specimens taken from a greenhouse in Singleton Park, Swansea, Glamorgan, 
in 1936, 1938 and 1949 ; and from a Reading University greenhouse at Shinfield, 
Reading, in 1959. Spirit specimens from Verdugo Woodlands, Glendale, California 
(collected by Dr. W. O. Gregg) ; Giiejar de la Sierra, Provincia de Granada, South 
Spain, and Elizondon, Navarre, North Spain (collected by Dr. de Zarate) and 
Barcelona. 



Litnax nyctelius Bourguignat, 1861 
Limax nyctelius Bourguignat, 1861 : 305, pi. 2, figs. 3 and 4. Algeria. 
First British record 

By Quick (1949 : 25). However, A. R. Waterston had found it some years before 
this, and partially described the species in an unpublished thesis. 

Diagnostic features 

Externally like Lehmannia poirieri, and with a similar radula. Internally resembles 
L. flavus in having a long rectal caecum, and a long cylindrical penis without an 
appendix. 

External appearance 

L. nyctelius is about 5 cm long, pale greyish-yellow with a thin translucent body- 
wall. There is a short inconspicuous keel at the hinder end. The mantle bands 
form a lyre-shaped figure, and the narrow body bands are high up on each side near 
the mid-dorsal line. The respiratory orifice has a pale border, and the sole is uniformly 
pale. 



BRITISH SLUGS 201 

External variation 

The mantle and body bands vary somewhat in breadth and intensity. Like those 
of Lehmannia poirieri, they tend to fade in alcohol. 

Jaw 

Dark brown or blackish, 17 mm across. 

Radula (Text-fig. 17 e) 

A tri-cuspid median tooth is flanked on each side by thirteen to fifteen tri-cuspid 
laterals and from twenty-eight to thirty aculeate marginals. The marginals beyond 
about the nineteenth are bifid, and the extreme marginals trifid. 

Alimentary system 

As in L. flavus, with two forwardly-directed loops and a long rectal caecum. 

Reproductive system (Text-fig. 17 b) 

The pigmented ovotestis is exposed between the right and left lobes of the liver. 
The pigmented hermaphrodite duct becomes swollen and folded in its central portion, 
and ends at the large albumen gland with a pale oval seminal vesicle. The free oviduct 
is short and stout. The spermatheca arises by a wide duct from the atrium, which 
expands abruptly to an oval sac which may be pigmented. The penis is uniformly 
cylindrical, more or less folded, and the vas deferens enters its apex under cover of 
the stout penial retractor muscle, which arises from the diaphragm close behind the 
kidney. Internally the penis (Text-fig. 17 c) resembles that of L. flavus in containing 
a prominent fold starting at the apex and expanding below to a " comb ", and 
another shorter smooth fold. 

Ecology 

Recorded from greenhouses and garden frames in Britain, on date palms in 
Washington, D.C., and beneath flower pots in Egypt (Abu Teira, in litt.). 

Distribution 

North African coast and Egypt. Introduced into Britain (known from greenhouses 
in Edinburgh and Glasgow), North America (known from Washington, D.C.) and 
South Africa (Connolly 1939 : 176). 

Material examined 

Spirit specimens from the Royal Botanic Gardens, Edinburgh (sent by A. R. 
Waterston), Shebin El Kom, Egypt, Algeria and Washington, D.C. 



202 BRITISH SLUGS 

FOSSIL SPECIES 

Limax modioliformis Sandberger, 1880 

Limax modioliformis Sandberger, 1880 : 103, pi. xii, figs. 15-15 c. 

Fossil shells ascribed to this species are found in the Cromerian at West Run ton, 
Norfolk (Sandberger, 1880). However, the specific ascription of fossil slug shells 
is at best only tentative. Similar shells in the same beds are ascribed to Lehmannia 
marginata. 





Fig. 18. Embryonic vesicles. A. Achatina fulica — embryo ( X 6-7) showing large 

caudal vesicle. B. A. fulica — later embryo (x 6-7) with vesicle adsorbed. 

G. Testacella scutulum — embryo ( X 6-7) showing cephalic and caudal vesicles. 

D. Helix aspersa — embryo (x 13) showing caudal vesicle, cephalic vesicle adsorbed. 

E. Helicella itala — embryo ( X 20) showing caudal vesicle, cephalic vesicle adsorbed. 

F. Avion ater — embryo (x 17) three weeks after laying, with cephalic and caudal 
vesicles. G. Agriolimax caruanae — embryo ( x 17) with cephalic and caudal vesicles. 

H. Limax cinereoniger — embryo ( x 6-7) with cephalic and caudal vesicles. 



BRITISH SLUGS 



203 



ANATOMICAL TERMS 

The following terms are used throughout this paper, in discussing the genital 
anatomy of slugs. They are illustrated by somewhat generalized illustrations in 
Text-fig. 19. 

The hermaphrodite gonad, far back in the visceral mass, producing 
ova and spermatozoa. 

The slender duct conveying ova and spermatozoa from the ovotestis 
to the albumen gland, oviduct and prostate. 

A loop, enlargement or diverticulum of the hermaphrodite duct 
at the albumen gland, for the storage of the animal's spermatozoa. 

A more or less linguiform gland at the termination of the herma- 
phrodite duct, supplying the egg albumen. 

A cavity at the commencement of the oviduct where the ova are 
fertilized by the partner's spermatozoa. 

Glandular tubules, opening into the male duct or groove which 
conveys the spermatozoa. 

The combined prostate, male duct or groove and oviduct. The 
prostate usually appears as a yellow ribbon on the gelatinous- 
looking oviduct. In the Stylommatophora, the proximal parts 
of the male and female tracts are not completely separate from 
one another. 

The slender male duct after it becomes separate and leaves the 
sperm-oviduct. 

The female duct, beyond the point where the vas deferens leaves it. 

The expanded termination of the spermatheca duct, which arises 
near the termination of the free oviduct (Milax) or directly from 
the atrium [Avion, Limax and Agriolimax). The spermatheca 
temporarily stores the partner's spermatozoa. 

The part of the free oviduct between the origin of the spermatheca 
duct and the atrium. When the spermatheca duct arises directly 
from the atrium, there is no vagina. 

The terminal portion of the genital ducts, receiving the oviduct, 
penis and often the spermatheca duct. 

The enlarged terminal portion of the vas deferens, which secretes 
the spermatophore and leads to the penis. 

A chitinous capsule containing a mass of spermatozoa [Avion, Milax). 

The terminal evaginable portion of the male duct, leading to the 
atrium, and sometimes (Milax) containing a penial papilla. 

A muscle arising from the body-wall and inserted between the 
epiphallus (when one is present) and the proximal end or apex 
of the penis. In Avion a penis and retractor are not present ; 
the epiphallus enters the atrium direct, and a retractor muscle 
is inserted on the free oviduct and the spermathecal duct. 

A slender tubular diverticulum from the apex of the penis (Testacella) . 

A conical or digitiform appendage of the penis (Lehmannia), or 
a more or less branched appendage at the apex of the penis 
(Agviolimax). 

A stimulatory organ situated in the penis (Agriolimax). 

A stimulatory organ situated in the atrium (Milax). 

A lobulated or frilled mass in the atrium or lower part of the oviduct 
in Avion, which in copula functions as an adhesive organ between 
the partners. 

A mass of glandular tubules in Milax, discharging into the termina- 
tion of the oviduct and the atrium. 



Ovotestis 



Hevmaphvodite duct 



Seminal vesicle 



Albumen gland 



Fevtilization sac 



Pvostate 



Spevm-oviduct 



Vas defevens 

Fvee oviduct 
Spermatheca 



Vagina 



A tvium 

Epiphallus 

Spevmatophove 
Penis . 

Retvactor penis muscle 



Flagellum 
Penial appendix 



Savcobelum 

Stimulatov 

Ligula 



Vaginal glands 



204 



BRITISH SLUGS 



A.Testacella 



s p — 





S P 



v D 



C. Milax 




VD 



D* Agriolimax 
S B 



H D 




Fig. 19. Genital anatomy. A. Testacella. B. Avion. G. Milax. D. Agriolimax. 



Key : A — atrium 

AG — -albumen gland 

E — epiphallus 

F — flagellum 
FO — free oviduct 
FS — fertilization sac 
HD — hermaphrodite duct 

L — ligula 
LA — lower atrium 
OT— ovotestis 

P — penis 
PA — penial appendix 



PR — prostate 

R — retractor muscle 
SB — sarcobelum 
SP — spermatheca 
SPD — spermatheca duct 
SPO— spermoviduct 
ST — stimulator 
SV — seminal vesicle 
UA — upper atrium 

V — vagina 
VD — vas deferens 
VG — vaginal glands. 



BRITISH SLUGS 



205 



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REFERENCES 

Adams, L. E. 1898. Observations on the pairing of Limax maximus L. /. Conch. 9 (3) : 

92-95 (pi. iii). 

19 10. Observations on the pairing of Avion ater (L.). Ibid. 13 (4) : 11 6-1 19 (6 text-rigs). 

1941. Testacella scutulum — method of cleansing. Ibid. 21 (8) : 223. 

Alder, J. 1848. A Catalogue of the mollusca of Northumberland and Durham. Trans. 

Tyneside Naturalists' Field Club 1 (2) : 97-209. 
Allman, G. J. 1843. On a new genus of terrestrial gastropod. Athenaeum (Sept.) : 851, col. 3. 

1846. Description of a new genus of pulmonary gasteropods. Ann. Mag. nat. Hist. 

17 : 297-299 (pi. xvii, A, figs. 1-3). 

Altena, C. O. van R. 1949. List of localities visited during a collecting trip to the Canary 
Islands in the spring of 1947- Tijdschr. Ent. 91 : 1-5. 

1950. The Limacidae of the Canary Islands. Zool. Verh. Leiden (11) : 3-34. 

1958. Notes sur les limaces 5. Les Philomycidae et Limacidae de la republique El Salvador. 

Arch. Molluskenk. 87 : 27-31 (2 figs.) 

Argenville, A. J. Dezallier d\ See Dezallier d'Argenville, 1742. 

Babor, J. F. 1894. liber den Cyclus der Geschlechtsentwicklung der Stylommatophoren. 
Verh. dtsch. zool. Ges. (4) : 55-61 (10 text-figs). 

Barnes, J. F. & Stokes, B. M. 1951. Marking and breeding Testacella slugs. Ann. appl. 
Biol. 38 (2) : 54°-545- 

Barnes, J. F. & Weil, J. W. 1945. Slugs in gardens : their numbers, activities and distribu- 
tion. Part 2. /. Anim. Ecol. 14 (2) : 71-105 (pis. 3-5). 

Baudon, A. 1884. Troisieme catalogue des mollusques vivants du departement de l'Oise. 
/. Conchyliol. 32 : 193-325. 

Binney, A. 1843. Descriptions of some of the species of naked, air-breathing mollusca, 
inhabiting the United States. Boston Journ. nat. Hist. 4 (2) : 163-175. 

1844. On some of the species of naked pneumonobranchus mollusca of the United States. 

Proc. Boston Soc. nat. Hist. 1 : 51-52. 

Binney, W. G. & Bland, T. 1869. Land and freshwater shells of North America. Pt. 1. 
Pulmonata Geophila. Smithson. misc. Coll. 8 (3) : i-xii -f- 1-3 16 (545 figs.). 

Bland, T. See Binney & Bland, 1869. 

Bonelli, — . In Lessona and Pollonera, 1882. 

Bouchard-Chantereaux, N. R. 1837. Catalogue des mollusques terrestres et fluviatile 
observes jusqu'a ce jour a l'etat vivant dans le departement du Pas-de-Calais. Mem. Soc. 
Agric. Boulogne (11) : 141-230. 

Bouillet, J. B. 1835. Catalogue des especes et varietes de mollusques terrestres et fluviatiles 
observes jusqu'a ce jour a 6tat vivant, dans la Haute et la Basse- Auvergne, etc. Ann. 
Scient. Acad. Sci. Clermont-Ferrand 8 : 521-594. 

Bourguignat, J. F. 1861. Des Limaces Algeriennes. Rev. Mag. Zool. (2) 13 : 299-306. (Re- 
printed in Spicileges Malac. pp. 33-43 with new continuation, 2 pp.) 

Boycott, A. E. 1934. Tne habitats of land mollusca in Britain. J. Ecol. 22 (1) : 1-38 (1 fig.). 

Breviere, L. 1 88 1. Tableau des limaciens des environs de Saint-Saulge (Nievre). /. Con- 
chyliol. 29 : 306-316. 

Broeck, E. V. 1870. Excursions decouvertes et observations malacologiques faites en Belgique 
pendant l'anee 1870. Ann. Soc. zool. malac. Belg. 5 : 13-64 (pi. 2 ; 1 text-fig.). 

Cain, A. J. & Williamson, M. H. 1958. Variation and specific limits in the Arion ater aggre- 
gate. Proc. malac. Soc. Lond. 33 (2) : 72-86 (5 figs.). 

Chemin, E. 1939^. Observations sur la Testacelle : Testacella europaea Roissy = T. haliotidea 
Drap. Trav. Sta. zool. Wimereux 13 : 77-85 (pi. 2 ; 1 fig.). 

19396. L'autofecondation chez Testacella europaea Roissy (= T. haliotidea Drap.). C.R. 

Soc. Biol. Paris 132 (24) : 249-252. 

zool. 6, 3. i5§ 



218 BRITISH SLUGS 

Clarke, B. J. 1843. On the species of the genus Limax occurring in Ireland. Ann. Mag. nat. 
Hist. 12 : 332-34 2 (3 Pis-)- 

In Thompson, 1840. 

Cockerell, T. D. A. 1 885. The variation and abnormal development of the mollusca. Part 
3. Terrestrial Gasteropoda. Hardwicke's Science Gossip 21 (250) : 224-226 (5 figs.). 

1886a. Notes. The Garner and Science Recorder's Journal 1 (9) : 139. 

18866. Notes on some varieties of British shells. /. Conch. 5 (3) : 79-80. 

1886c. Some varieties of British shells. The Naturalist's World 3 (33) : 179. 

1890a. The British naturalist's catalogue of the land and freshwater mollusca of the 

British Islands. The Young Naturalist 11 (127) : 19 pp. 

18906. Notes on slugs, chiefly in the collection of the British Museum. Ann. Mag. nat. 

Hist. (6) 6 : 277-288. 

1 891. Notes on slugs, chiefly in the collection of the British Museum. Ibid. (6) 7 : 328-341. 

— — 1892. A revised list of the species of British Slugs. /. Conch. 7 (3) : 66-70. 

Collinge, W. E. 1892a. Descriptions of a new variety of Arion hortensis Fer. and A. circum- 
scripta Johnst. The Conchologist 2 (2) : 26-27. 

18926. A review of the Arionidae of the British Isles. Ibid. 2 (3) : 56-66. 

1892c. A review of the Arionidae of the British Isles. Ibid. 2 (4) : 76-83. 

1893. On the occurrence of Arion lusitanicus Mab. in the British Isles, and descriptions 

of four new varieties. Ann. Mag. nat. Hist. (6) 12 : 414-415. 

1894. The anatomy and description of a new species of Arion. Ibid. (6) 13 : 66-67 (p*- 

V.A.). 

1895a. Amalia parryi, a supposed new species. Journ. Malac. 4 (1) : 7. 

■ 18956. Notes on some slugs from Algiers. Proc. malac. Soc. Lond. 1 (7) : 336-337 (pi. 

xxiii) . 

1896. On a collection of slugs from the Sandwich Islands. Ibid. 2 (1) : 46-51 (6 figs.). 

1897a. Some observations on certain species of Arion. Journ. Malac. 6 (1) : 7-10 (pi. 2). 

18976. On some European slugs of the genus Arion. Proc. zool. Soc. Lond. (3) : 439-450 

(pis. xxix-xxxi). 
1897c. On a further collection of slugs from the Hawaiian (or Sandwich) Islands. Proc. 

malac. Soc. Lond. 2 (6) : 293-297 (9 figs.). 

■ 1898. Note on a new variety of Testacella maugei Fer. /. Conch. 9 (3) : 95. 

Comte, T. le. 1871. Deuxieme liste supplementaire des mollusques terrestres et fluviatiles 

recueilles aux environs de Lessines. Bull. Soc. mat. Belg. 6 : 65—68. 
Conchological Society Census. See Ellis, 195 1. 
Connolly, M. 1939. A monographic survey of South African non-marine mollusca. Ann. 

S. Afr. Mus. 33 (1) : i-iii + 1-660 (pis. I-XIX ; 58 text-figs.). 
Cooper, J. G. 1863. On a new genus of terrestrial mollusca inhabiting California. Proc. Calif. 

Acad. Sci. 3 (1) : 62-63 i 1 n g-)- 
Dezalier d'Argenville, A. J. 1742. L'Histoire Naturelle eclaircie dans deux de ses parties 

principales, la lithologie et la conchyliologie . . . viii + 491 pp. (33 pis.) 4T.0. Paris. 
d'Orbigny, A. D. See Orbigny, A. D. d', 1836-44. 
Draparnaud, J. P. R. 1801. Tableau des mollusques terrestres et fluviatiles de la France. 

116 pp. 8vo. Montpellier & Paris. 
1805. Histoire naturelle des mollusques terrestres et fluviatiles de la France, i-viii + 1-164 

pp. (13 pis. 1 port.) 4to. Paris. 
Drouet, H. 1868. Mollusques terrestres et fluviatiles de la Cote-d'Or. Mem. Acad. Dijon. 

(2) 14 : 33-154. 
Druce, G. C. 1932. The comital flora of the British Isles, xxxii + 407 pp. Arbroath. 
Dumont, F. 1850. Description d'une espece nouvelle du genre Arion (A. lineatus). Bull. 

Soc. Hist. nat. Savoie 1 : 64-65. 
Dumont, F. & Mortillet, G. 1 856. Catalogue critique et malacostatique des mollusques 

de Savoie et du bassin du Leman. Bull. Inst. nat. Genevois 4 : 310-361. 



BRITISH SLUGS 219 

Ellis, A. E. 1926. British Snails. A guide to the non-marine gastropoda of Great Britain and 

Ireland Pliocene to Recent. 275 pp. (14 pis., 11 text-figs.). Clarendon Press, Oxford. 

1941. The mollusca of a Norfolk Broad. /. Conch. 21 (8) : 224-243. 

1951- Census of the distribution of British non-marine mollusca. Ibid. 23 (6 and 7) : 

171-244 (8 pp. maps). 
Esmark, B. 1883. Nyt Bidrag til Kundekaben om Norges Land og Ferskvands-Mollusker. 

Nyt. Mag. Naturv. 27 : 77-110. 
Ferussac, J. B. L. d'A. de. 1819. Histoire naturelle generate et particuliere des mollusques 

terrestres et fluviatiles 2 : i-xvi + 1-96. Fol. Paris. 

1823. Op. cit. (Nouvelle division des pulmonis sans opercule) 2 : 96 a-z, a-A. 

Fischer, P. H. See Gassies & Fischer, 1856. 
Gaimard, J. P. See Quoy & Gaimard, 1824. 
Gassies, J. B. 1849. Table methodique et descriptif des mollusques terrestres et d'eau douce de 

VArgenais. 209 [4] pp. (4 pis. col.). 8vo. Paris. 
Gassies, J. B. & Fischer, P. H. 1856. Monographic du genre Testacelle. Act. Soc. linn. 

Bordeaux 21 : 195-248 (2 pis.). 
Gerhardt, U. 1933. Zur Kopulation der Limaciden. Z. Morph. Okol. Tiere 27 (3) : 401-450 

(11 text-figs.). 
1935- Weitere Untersuchungen zur Kopulation der Nacktschnecken. Ibid. 30 (2) : 

297-332 (9 ngs.). 

1939. Neue biologische Untersuchungen an Limaciden. Ibid. 35 (2) : 183-202 (3 figs.). 

1940. Neue biologische Nacktschneckenstudien. Ibid. 36 (4) : 557-580 (8 figs.). 

Germain, L. 1930. Mollusques terrestres et fluviatiles. Faune Fr. 21 : 447 pp. (13 pis. 470 

figs-)- 

Gistel, J. von N. F. X. 1848. Naturgeschichte des Thierreichs fur hohere Schulen. xvi + 216 
[4] pp. (32 pis. col.). Fol. Stuttgart. (Mollusca pp. 165-174.) 

Gray, J. E. 1821. A natural arrangement of mollusca, according to their internal structure. 
London Medical Repository 15 : 229-239. 

— — ■ 1824. On the natural arrangement of the pulmonobranchous mollusca. Annals of Philo- 
sophy, Fond, (n.s.) 24 : 107-109. 

1840. A manual of the land and freshwater shells of the British Islands, with figures of each 

of the kinds. By William Turton (A new edition thoroughly revised and much enlarged by 
J. E. Gray) ix [i] + 324 pp. (12 pis. col., text illustrated). 8vo. London, 1 Apr. 

1855. Catalogue of Pulmonata or air-breathing mollusca in the collection of the British 

Museum. Part 1. [iv] + 192 pp. (text illustrated). 12 mo. London, 10-18 Apr. 

Gregg, W. O. 1944. Proterogyny in Deroceras panormitanum. Nautilus 58 : 67-68. 
Hayward, J. F. 1954a. Agriolimax caruanae Pollonera as a Holocene fossil. /. Conch. 
23 (12) 1403-404 (pi. 15). 

I954&- A further note on Agriolimax caruanae Pollonera. Ibid. 24 (1) : 21-22. 

Hazay, J. 1 881. Die Molluskenfauna von Budapest. Malakozool. Bl. (n.s.) 3 : 1-69 (pi. 1, 

figs. 1, 2). 
Held, F. 1836. Aufzahlung der in Bayern lebenden Mollusken. Isis 29 (4) : 271-282. 
1848-49. Die Landmollusken Bayerns. Jahresb. der K. Kreis-Landwirtschafts- und Gewerbs- 

Schule zu Munchen fur das Schuljahr, i848~4g : 23. 4to. Miinchen. 
Hesse, P. 1926. Die Nacktschnecken der palaearktischen Region. Abh. Arch. Molluskenk. 

2 (1) : 1-152 (2 pis.). Frankfurt. 
Heynemann, D. F. 1861. Die nackten Schnecken des Frankfurter Gebiets, vornehmlich 

aus der Gattung Limax. Malakozool. Bl. 8 : 85-105. 

1873. Ueber Geomalacus. Ibid. 21 : 25-36 (taf. 1). 

See Koch & Heynemann, 1874. 

Hoffmann, H. 1925. Zur Kenntnis der Testacellen. Arch. Molluskenk. 57 (5 and 6) : 171- 

221 (taf. v, vi). 
Hutton, F. W. 1879. Description of some new slugs. Trans, Proc, N,Z. Inst. 11 : 331-332. 



220 BRITISH SLUGS 

Jeffreys, J. G. 1862. British Conchology, Vol. I, land and freshwater shells, cxiv + 341 pp. 

(8 pis.). London. 
Jensen, O. S. 1873. Indberetning om en i Sommeren 1870 foretagen Reise i Kristiania og 

Kristianssands Stift forat undersoge Land og Ferskvands-Molluskerne tilligemed Iglerne. 

Nyt. Mag. Naturv. 19 : 146-188 (pi. 1). 
Johnston, G. 1828. A few remarks on the class mollusca, in Dr. Fleming's works on British 

animals ; with descriptions of some new species. Edinburgh new Phil. Journ. 5 : 74-81. 

1838. A list of the pulmoniferous mollusca of Berwickshire and North Durham. Berwick- 
shire Naturalists' Club 1 : 154-156. 

Kaleniczenko, J. 1839. Necrologie : Notice sur le Professor J. A. de Krynicki. Bull. 

Soc. Imp. Nat. Moscou (1) : 25-33. 
1 85 1. Description d'un nouveau genre de limaces de la Russie meridionale. Ibid. 24 : 

215-228 (pis. 5, 6). 
Kennard, A. S. & Woodward, B. B. 1926. Synonymy of the British non-marine mollusca 

{Recent and Post-tertiary), xxiv -\- 447 pp. London. 
Kew, W. 1902. On the mucous-threads of land-slugs. /. Conch. 10 (5) : 153-165. 
Koch, C. & Heynemann, D. F. 1874. Neue Nacktschnecken aus Turkestan. Jb. dtsch. 

malakol. Ges. 1 : 1 51-153. 
Kunkel, K. 1916. Zur Biologic der Lungenschnecken Ergebnisse vieljdhriger Zuchtungen 

und Experimente. xvi + 440 pp. (48 text-figs., col. pi.). Heidelberg. 
le Comte, T. See Comte, T. le. 

Lehmann, R. 1864. Neue Nacktschnecke aus Australien. Malakozool. Bl. 11 : 145-149. 
Lessona, M. 1880. Molluschi viventi del Piemonte. Mem. Accad. Lincei (3) 7 : 317-380 (4 pis.). 
Lessona, M. & Pollonera, C. 1882. Monografia dei Limacidi Italiani. 73 pp. (3 pis.). 4to. 

Torino (preprint of Mem. R. Accad. Torino (2) 35 : 49-128, 1884). 
Leydig, F. 1876. Die Hautdacke und Schale der Gastropoden, nebst einer Uebersicht der 

einheimischen Limacinen. Arch. Naturgesch. 42 (1) : 209-292 (pis. ix-xvi). 
Linnaeus, C. 1758. Sy sterna Naturae . . . Editio decima, reformata 1 [iv] -f 824 pp. 8vo. 

Holmiae. 

1767. Sy sterna Naturae . . . Editio duodecima reformata 1 (2) : 533-1327. 8vo. Holmiae. 

Lister, M. 1674. An extract of a letter of Mr. Martin Lister concerning the first part of his 

tables of snails, together with some queries relating to those Insecta, and the tables themselves. 

(Some general queries concerning land and freshwater Snails.) Philosophical Transactions, 

9 (105) : 96-99 (pi. 2). 

1685-97. Historiae sive synopsis methodicae conchyliorum, etc. 6 pts. Fol. Londini, 1685- 

92 (-97). 
Locard, A. 1895. Ipsa Draparnaudi conchylia. Iitude sur la collection conchyliologique de 

Draparnaud au Musee Imperial et Royal d'Histoire Naturelle de Viennes. 190 pp. Paris. 
Lohmander, H. 1937- Uber die nordischen Formen von Arion circumscriptus Johnston. 

Acta. Soc. Fauna Flora fenn. 60 : 90-112 (17 figs.). 
Lukis, F. C. 1834. A notification of the occurrence, in the Island of Guernsey, of a species of 

Testacellus, and of some of its characteristics and habits, as observed there. Mag. nat. Hist. 

7 : 224-226 (2 figs.). 
Luther, A. 1915. Zuchtversuche an Ackerschnecken (Agriolimax reticulatus Mull, und 

Agr. agrestis L.). Acta. Soc. Fauna Flora fenn. 40 (2) : 1-42 (2 text-figs.). 
Mabille, M. J. 1867. Archives malacologiques ... I. Le genre Geomalacus. Rev. Mag. 

Zool. (2) 19 : 53-64. 

1868. Des limaciens europeens. Rev. Mag. Zool. (2) 20 : 129-146. 

1883. Sur quelques especes de mollusques terrestres. Bull. Soc. philom. Paris (7) 7 : 

39-53- 
Makings, P. 1959. Agriolimax caruanae Pollonera new to Ireland. /. Conch. 24 (10) : 354-356. 
Malm, A. W. 1868. Zoologiska observationer. V. Hdftet. Skandinaviska Land-Sniglar, Lima- 

cina etc. (5 pis.). 8vo. Goteborg. (A preprint from Goteborgs VetenskSamh. Handl, 1870, 

10 : 26-93.) 



BRITISH SLUGS 221 

Millet, P. A. 1843. Description de plusieurs especes nouvelles de mollusques de France. 

Mag. Zool. Paris. (2) (Classe V) Mollusques, 4 pp. (pis. 63, 64). 
Moquin-Tandon, A. 1855-56. Histoire naturelle des mollusques terrestres et fluviatiles de 

France, etc. 2 torn, (illust.). 8vo. Paris. 
Morch, O. A. L. 1865. Quelques mots sur un arrangement des mollusques pulmon6s terrestres 

(Geophiles, Fer.) bas6 sur le systeme naturel (suite). /. Conchyliol. 13 : 376-396. 
Morelet, P. M. A. 1845. Description des Mollusques terrestres et fluviatiles du Portugal. 

8vo. Paris (July). 
Mortillet, G. See Dumont & Mortillet, 1856. 
Muller, O. F. 1774. Vermium terrestrium et fluviatilium . . . Historia. 2. xxxvi -f- 214 -f- [10]. 

4to. Havniae & Lipsiae. 
Nilsson, S. 1822. Historia molluscorum sveciae terrestrium et fluviatilium breviter delineata. 

xx -f- 124 pp. 8vo. Lundae (1822, 1823). 
Normand, N. A. J. 1852. Description de six limaces nouvelles observees aux environs de 

Valenciennes. 8 pp. 8vo. Valenciennes, Nov. 
0kland, F. 1923. Arionidae of Norway. Skr. VidenskSelsk., Christ. 1922 (5) : 1-62 (pi. 1, 

46 figs., 5 maps). 
Oldham, C. 1942a. Auto-fecundation and duration of life in Limax cinereo-niger. Proc. malac. 

Soc. Lond. 25 (1) : 9-10. 

19426. Notes on Geomalacus maculosus. Ibid. 25 (1) : 10-11. 

Orbigny, A. C. V. M. d'. 1836-44. Mollusques. Histoire Naturelle des lies Canaries. (Webb & 

Berthelot) 2 (2) : 4-1 17 (pis. i-vii B). 
Philippi, R. A. See Schultz, A. W. F. 1836-44. 
Phillips, R. A. & Watson, H. 1930. Milax gracilis (Leydig) in the British Isles. J. Conch. 

19 (3) : 65-93 (pis. 1, 2). 
Pilsbry, H. A. 1944. West American field slugs (Deroceras). Nautilus 58 : 15-16. 

1948. Land Mollusca of North America (North of Mexico). Monogr. Acad. nat. Sci. 

Philad. (No. 3) : 2 i-xlvii + 521-1113 (figs. 282-585). 

Pini, N. 1876. Molluschi terrestri e d'acqua dolce viventi nel territorio d'Esino. Bull. Soc. 

Malac. Ital. 2 : 67-205 (2 pis.). 
Plate, L. H. 1891. Studien uber opisthopneumone Lungenschnecken. Die Anatomie der 

Gattungen Daudebardia und Testacella. Zool. Jb. 4 : 505-630 (pis. 32-37). 
Pollonera, C. 1887a. Intorno ad alcuni Limacidi europei poco noti. Boll. Mus. Zool. 

Anat. comp. Torino 2 (21) 14 pp. (1 pi). 
18876. Specie nuove o mal conosciute di Arion europei. Atti Accad. Torino 22 (5) : 290- 

313 (1 pl)- 

1889. Nuove contribuzioni alio studio degli Arion europei. Ibid. 24 : 623-640 (1 pl.). 

1891. Appunti di Malacologia. Intorno ai Limacidi di Malta. Boll. Mus. Zool. Anat. 

comp. Torino 6 (99) : 1-4 (2 figs.). 

1882. See Lessona & Pollonera, 1882. 

Quick, H. E. 1946. The mating process in Arion hortensis Ferussac and in Arion subfuscus 

Draparnaud. /. Conch. 22 (8) : 178-182 (8 figs.). 
1947- Arion ater (L.) and A. rufus (L.) in Britain and their specific differences. Ibid. 22 

(10) : 249-261 (23 figs.). 
— — 1949. Synopses of the British fauna. No. 8. Slugs {Mollusca). (Testacellidae, Arionidae, 

Limacidae.). 29 pp. (2 pis. 26 text-figs.). Linnean Society of London. 

1950- The spermatophore of Milax sowerbii (Ferussac). /. Conch. 23 (4) : 111-112 (8 

figs.). 

1952. Rediscovery of Arion lusitanicus Mabille in Britain. Proc. malac. Soc. Lond. 29 

(2 and 3) : 93-101 (15 fi gs.). 

1954. Journey to the Gugh6 Highlands (Southern Ethiopia). 1948-49 : Gastropoda, 

slugs from the high mountains. /. Linn. Soc. {Zool.) 42 : 382-386 (17 figs.). 

Quoy, J. R. C. & Gaimard, J. P. 1824. Voyage autour de monde . . . execute sur . . . 
I'Uranie et la Physicienne . . . Zoologie text. 671 pp. 4T.0. Paris. 



222 BRITISH SLUGS 

Rafinesque-Schmaltz, C. S. 1815. Analyse de la nature, etc. 8vo. Palerme. 

1820a. Annals of Nature or Annual Synopsis of new genera and species of animals, plants 

etc. discovered in North America . . . First Annual Number, for 1820. 16 pp. 8vo. Lex- 
ington. 
18206. Monographie des coquilles bivalves fluviatiles de la Riviere Ohio, etc. Ann. 

gen. Sci. phys. 5 : 287-322 (3 pis.). 
Razoumowsky, G. de. 1789. Histoire naturelle du Jorat et de ses environs, etc. 1. 8vo. 

Lausanne. (Mollusca pp. 266-278.) 
Recluz, C. A. 1845. Description des Mollusques terrestres et fluviatiles du Portugal, par A. 

Morelet. Rev. zool. 1845 : 306-312. 
Risso, A. 1826. Histoire naturelle des principales productions de V Europe meridionale, et 

particulierement de celles des environs de Nice et des Alpes Maritimes. 4. 8vo. Paris. 
Roebuck, W. D. 1883. The British Slug List. /. Conch. 4 (2) : 38-43. 

■ 1884a. A new variety of Arion ater. Ibid. 4 (5) : 146. 

18846. Arion ater var. bicolor in West Gloucestershire. Ibid. 4 (7) : 217. 

1885. New varieties of Limax arborum and Arion ater. Ibid. 4 (12) : 375-376. 

— — 192 1. Census of the distribution of British land and freshwater mollusca. Ibid. 16 (6) : 

165-212. 
Roissy, A. F. P. M. de. 1805. Histoire naturelle . . . des Mollusques, etc. (Suites a 

Hist. Nat. Buffon. redig. Sonnini.) 5 and 6. 8vo. Paris, An. 13. 
Sandberger, C. L. F. von. 1880. Ein Beitrag zur Kenntnis der unterpleistocanen Schichten 

Englands. Palaeontographica (n.f. 7) 27 (2) : 83-104 (1 pi.). 
Schrank, F. von P. 1803. Fauna Boica. 8vo. Landshut. (Preface dated 8 Nov., 1802.) 
Schrenk, . 1848. Uebersicht der Land und Susswasser — Mollusken Livlands. Bull. Soc. 

imp. Nats. Moscou 21 (1) : 135-185. 
Schultz, A. W. F. 1836-44. in Philippi, R. A. Enumeratio Molluscorum Siciliae, etc. 2 vols. 

4to. Berolini, etc. 
Siebert, H. 1873. Die colorirten Tafeln des Lehmann' schen Werkes. NachrBl. dtsch. 

malakozool. Ges. (now Arch. Molluskenk.) 5 : 79-82. 
Simroth, H. 1885. Versuch einer Naturgeschichte der Deutschen Nacktschnecken und 

ihrer Europaischen Verwandten. Z. wiss. Zool. 42 (2) : 203-366 (pis. 7-1 1). 
1887. Einige Bemerkungen, betreffend die Systematik der europaischen Nacktschnecken. 

NachrBl. dtsch. malakozool. Ges. (now Arch. Molluskenk.) 19 : 161-168. 
1906. Ueber eine Reibe von Nacktschnecken die Herr Dr. Cecconi auf Cypern und in 

Palaestina gesammelt hat. Ibid. 38 (2) : 84-91. 
Sordelli, F. 1870. Anatomia del Limax Doriae, Bourg., etc. (Descrizione succinta delle due 

species di Limax etc.). Atti Soc. ital. Sci. nat. 31 : 242-253 (1 pi.). 
Sowerby, G. B. 1820. The genera of recent and fossil shells . . . continued by G. B. Sowerby. 

pt. 1. (first issue), pages not numbered. 8vo. London. 
Stokes, B. M. 1958. The worm-eating slugs Testacella scutulum Sowerby and T. haliotidea 

Draparnaud in captivity. Proc. malac. Soc. Lond. 33 (1) : 11-20. 
— — See Barnes & Stokes, 1951. 
Taylor, J. W. 1902-07. Monograph of the land and freshwater mollusca of the British Isles. 

(Testacellidae, Limacidae, Arionidae). (pts. 8-13) : i-xx + 1-312 (25 pis. 275 figs, portraits). 

Leeds. 
Thomas, D. C. 1944. Field sampling for slugs. Ann. appl. Biol. 31 (2) : 163-164. 
Thompson, W. 1840. Catalogue of the land and freshwater mollusca of Ireland. Ann. Mag. 

nat. Hist. 6 : 194-206. 
Turton, W. 1831. A manual of the land and freshwater shells of the British Islands, viii + 

150 -f- [16] pp. (10 pis., col.). 8vo. London. 

See Gray, 1840. 

Watson, H. 191 5. Studies on the carnivorous slugs of South Africa, including a monograph 

of the genus Apera, and a discussion on the phylogeny of the Aperidae, Testacellidae, and 

other agnathous Pulmonata, Ann. Natal Mus. 3 (2) : 107-267 (pis. 7-24 ; 7 text figs.). 



BRITISH SLUGS 



223 



Watson, H. 1930. Milax gracilis (Leydig) in the British Isles. Part 2. Anatomy. J . Conch. 

19 (3) : 72-93 (pis. 1 and 2, text-figs. 8-23, map). 

1943- Notes on a list of the British non-marine Mollusca. Ibid. 22 (3) : 53-72. 

See Phillips and Watson, 1930. 

Webb & Berthelot. See Orbigny, 1836-44. 

Weil, J. W. See Barnes & Weil, 1945. 

Weinland, D. F. 1876. Zur Wiechthierfauna der Schwabischen Alb. Jhr. Ver. vaterl. Naturk. 

Wurttemb. 32 : 234-358 (taf. 4). 
Westerlund, C. A. 1876. Neue Binnenmollusken aus Sibirien. NachrBl. dtsch. malakozool. 

Ges. (now Arch. Molluskenk.) 8 (8 and 9) : 97-104. 
Williams, J. W. 1888. The Shell-collector's handbook for the field, i-xii + 1-148 (14 figs.). 

8vo. London. 

See Cain & Williamson, 1958. 
Die Wiirmer. Sturm's Deutschlands Fauna, etc. (Abth. 6. Hft. 1 and 2). 



Williamson, M. H. 
Wolf, J. 1803-06. 
8vo. 

WOLLASTON, T. V. 

Woodward, B. B. 



1878. Testacea Atlantica, etc. pp. 
See Kennard & Woodward, 1926. 



xi -f- 588. 8vo. London. 



224 



BRITISH SLUGS 



INDEX 
Page 
agrestis, Agriolimax 163-4, ID 7> 168-70, 178-9 



agrestis, Limax 
Agriolimax 132, 



144, 163-80, 181 ; 



163, 
I94> 



168 



alba, Avion ater (var.) 
alba, Avion intermedins (var.) 
alba, Avion subfuscus (var.) 
alba, Milax sowerbyi (var.) 
albida, Arion atev (var.) 
albina, Testacella haliotidea (var.) 
albina, Testacella maugei (var.) . 
albolateralis, Arion ater (var.) 
aldvovandi, Limax maximus (var.) 
allmani, Geomalacus maculosus (var.) 
Amalia ..... 
Ambigolimax .... 
andvewsi, Geomalacus maculosus (var 
antiquovum, Limax . 
antiquorum, Limax fiavus (var. 
aperta, Testacella maugei (var.) 
avbovum, Limax 
A vion 



120, 124-47, 153, 



Arionidae 
Arioninae 
atev, Avion 



124, 126, 128, 



1 3o-3 1: 
1 36-: 



atev, Avion atev 

atev, Avion atev (var.) 

atev, Limax .... 

atevvima, Avion atev (var.) . 

auvea, Testacella maugei (var.) 

auvea, Testacella scutulum (var.) 

auvigastev, Testacella 

babovi, Amalia 

benoiti, Milax g agates (var.) 

bettonii, Lehmannia marginata (var.) 

bevenoti, Agviolimax 

bicolov, Avion subfuscus (var.) 

bicolov, Milax gagates (var.) 

bilobatus, Limax 

Binneya ..... 

bisulcata, Testacella . 

bveckwovthiana, Limax fiavus (var.) 

bvunnea, Avion intevmedius (var.) 

bvunneus, Limax 

budapestensis, Amalia 

budapestensis, Milax 149, 155, 

caevulea, Avion hovtensis (var.) . 
campestvis , Limax 
canaviensis, Limax 



203 
139 
125 
133 
158 
139 
115 
in 

139 
192 
120 
50 
194 
120 
191 
. 184 
in 

• 194 

183, 203 

119-47 

119-47 

133-34. 

, 138-47 

138-45 

. 139 



149- 



187, 



124 
139 



149- 



172 



138 

145 
in 
117 
no 
50 

151 
194 

175 

• 133 

• 151 
170 

. 119 
. 118 
. 184 

• 125 
172 
160 

158-60, 

160-63 

. 130 

172 

164 



Page 
Candida, Limax maximus (var.) . .192 

Cavinavion . . . . 124, 128 

cavinatus, Limax . . . .158 

cavuanae, Agviolimax 164,167, 169,173, 175-80 
castanea, Avion atev (var.) . . .139 

cellavia, Limax maximus (var.) . .192 

cevea, Limax tenellus (var.) . . 181 

ceveus, Limax . . . . .180 

cincta, Limax tenellus (var.) . . 181 

cinevacea, Agviolimax veticulatus (var.) 165 
cinevea, Limax cineveonigev (var.) . 188 

cineveus, Limax . . . .191 

cineveofusca, Avion subfuscus (var.) . 133 
cineveonigev, Limax 180, 183, 187-91, 192-3 
civcumscvipta, Avion fasciatas (var.) . 128 
civcumscviptus, Avion . . .127 

coccinea, Avion atev (var.) . . .139 

coevuleus, Avion . . . .130 

collingei, Amalia . . . .158 

concolov, Limax maximus (var.) . . 192 

cypvius, Agviolimax . . . .164 

Devocevas . . . . 163, 179 

elongatus, Avion . . . .130 

empivicovum, Avion . . 124, 138, 145 

Endodontacea .... 119-47 

Endodontidae . . . . .107 

euvopea, Testacella . . . 115 

fasciata, Geomalacus maculosus (var.) . 120 
fasciata, Limax maximus (var.) . . 192 

fasciatus, Avion 124-5, 127-9, 130-4, 144 
fasciatus, Limax . . . .127 

flavescens, Avion fasciatus (var.) . . 128 

flavescens, Testacella haliotidea (var.) . 115 
flavogviseus , Avion lusitanicus (var.) . 136 
fiavus, Avion . . . . .125 

fiavus, Limax 180, 184-7, 190, 196, 198, 200-1 
fuliginea, Avion subfuscus (var.) . 133 

fulva, Limax tenellus (var.) . . 181 

fulvus, Limax . . . . .180 

fuscatus, Avion . . . 133 

fuscocavinata, Milax sowevbyi (var.) . 158 
gagates, Limax . . . 149-50 



gagates, Milax 
Geomalacus 



149-55, 156-61 
119-24, 137 



glauca, Lehmannia mavginata (var.) 

globosus, Agviolimax . 

gvacilis, Limax 

gvisea, Agviolimax laevis (var.) 

gvisea, Avion fasciatus (var.) 

gvisea, Avion hovtensis (var.) 

gviseonigvescens, Testacella maugei (var. 



194 

175 
160 

173 
138 
140 
in 



BRITISH SLUGS 



225 



griseorubescens , Testacella maugei (var.) 11 1 
gughensis, Agriolimax . . .164 

haliotidea, Testacella 108, 109-10, in, 114, 

115-6, 1 1 7-8 
hiberna, Avion ater (var.) . .139 

hortensis, Avion 124, 128-9, 130-2, 133-4, 

I37> 144 
Hydrolimax . . . . .164 

hyperboreus , Limax . . . .172 

ichnusae, Amalia . . . .156 

insularis, Amalia . . . .156 

insularis, Milax . . 150, 155, 156-7 

intermedia, Limax cinereus (var.) 
intermedins, Avion 124, 125-7, 



Kobeltia ..... 
Kvynickillus .... 
lacustvis, Agviolimax laevis (var.) 
laevis, Agviolimax 164, 167, 172-5 
laevis, Limax 
Lehmannia 
Limacidae 
Limacinae 
Limacus 

Limax . 178, 180-93, 194, 1 
lineatus, Avion 
Lochea .... 
lusitanicus, Avion 124, 133, 135-8, 144-5 

maculata, Agviolimax laevis (var.) . 173 

maculata, Amalia . . . .158 

maculosus, Geomalacus . 119, 120-4, 137 
majov, Testacella haliotidea (var.) . 115 

mavginata, Lehmannia 193, 194-7, 198-200, 

202 
mavginatus, Limax . . .158, 193-4 



. 187 

129, 131, 

134, 144 

. 124 

164 

• 173 

178-9 

172 

193-200, 203 

147-202 

163-202 

. 180 

), 200-2, 203 

. 187 

. 124, 133 



mavginatus, Milax . 
mavginella, Avion atev (var.) 
maugei, Testacella . . 109-14 

mauva, Limax cineveonigev (var.) 
mauvus, Limax .... 

maximus, Limax 180, 183, 187-90, 191-3 
media, Avion atev (var.) 
meditevvanea, Amalia 
megalodontes , Limax 
melitensis, Lehmannia 
Mesavion .... 

Micvoavion . . . . .124 

Milax . 149-63, 165, 178, 181, 183, 203 
minimus, Avion . . . .125 

minutus, Kvynickillus . . .164 

modiolifovmis , Limax . . . 202 

nigva, Agviolimax veticulatus (var.) . 165 
nigva, Avion hovtensis (var.) . .130 



. 149 

• 139 

115-18 

. 188 

150 

197 

139 

150 

184 

193-4 

124 



nigva,' Testacella maugei (var.) 

nigvescens , Avion lusitanicus (var 

nigvescens , Milax sowevbyi (var.) 

nigvicans, Avion subfuscus (var.) 

nigvicans, Pavmacella 

nobvei, Avion . 

novmalis, Avion intevmedius (var 

nyctelius, Limax 

Oleacinacea 

Oleacinidae 

olivaceus, Avion lusitanicus (var. 

ovetea, Milax sowevbyi (var.) 

ovnata, Limax cineveonigev (var.) 

pallidus, Limax 

panovmitanus, Devocevas 

panovmitanus , Agviolimax . 

Pavmacella 

Parmacellinae 

pavvyi, Amalia 

pegovavii, Avion subfuscus (var.) 

pellucida, Vitvina 

pevkinsi, Agviolimax 

plumbea, Avion atev (var.) . 

plumbea, Avion intevmedius (var. 

plumbea, Milax gagates (var.) 

poivievi, Lehmannia 

poivievi, Limax 

punctata, Limax cineveonigev (var 

vava, Milax gagates (var.) . 

reticulata, Agviolimax veticulatus 

veticulata, Avion atev (var.) 

veticulatus, Agviolimax 163, 164-70 



in 

■ 136 
. 158 

• 133 
. 150 

• 135 

• 125 
180, 200-1 

108-18 
107 

• 136 
160 

. 188 
170 
. 179 
164 
. 149 
149-63 
149-50 

• 133 



158 



(var.) 



veticulatus, Limax 

vosea, Lehmannia mavginata (var.) 

vufescens, Agviolimax veticulatus (var.) 

vufescens, Avion hortensis (var.) 

rufescens, Arion lusitanicus (var.) 

rufescens, Limax flavus (var.) 

rufofusca, Arion subfuscus (var.) 

vufus, Avion atev . 138, 139, 144, 

vufus, Limax .... 

vupicola, Lehmannia mavginata (var.) 

vusticus, Milax 

vutilans, Geomalacus maculosus (var.) 

scotti, Agviolimax 

scutulum, Testacella 108-10, 111, 114, 



124 
175 
139 
125 
151 
193-4, 197-200, 201 
197 

, 188 
151 
165 
139 

171-2, 

177-9 

164 

194 

165 
130 

136 
184 

133 
145-7 
145 
194 
149 
120 
164 
116, 



sevotinus, Limax 

similis, Amalia meditevvanea 

sowevbyi, Limax 

sowevbyi, Milax . 149- 

subflavus, Avion 



50. 



117-8 

. 180 
• ) • 150 

• 158 
155, 158-60 

• *33 



226 



BRITISH SLUGS 



subfuscus, Avion 



124, 129, 133-5 



subfuscus, Limax 

subrufa, Lehmannia marginata (var.) 

Succinea ..... 

succinea, Avion atev (var.) . 

succinea, Avion subfuscus (var.) 

sulcatus, Avion 

sylvatica, Limax maximus (var.) 

tenellus, Agviolimax . 

tenellus, Limax 180-4, 185-6, 1 

Testacella . . . 108- 

Testacellidae .... 

tetvazona, Limax maximus (var.) 



Page 

136-7, 

44, 147 

133 

194 

109 

139 

133 

145 

192 

180 

38, 190 

18, 203 

108-18 

192 



is, Limax maximus (var.) 
tigrina, Lehmannia mavginata (var.) 
tvigona, Testacella haliotidea (var.) 
valentianus, Limax . 
vaviegatus, Limax . 
verkvuzeni, Geomalacus maculosus (var.) 
vinosa, Limax maximus (var.) 
violacea, Agviolimax veticulatus (var.) . 
vividans, Testacella maugei (var.) 
vivescens, Limax flavus (var.) 
Vitrina .... 
Zonitacea 
Zonitidae 
Zonitoides 



Page 
192 
194 
115 
197-8 
184 
120 
192 

• 165 
in 

. 184 

. i 7 8 

147-202 

107 

• 174 



I. Geomalacus maculosus. 

4. Avion hovtensis. 
6. Avion intevmedius. 



9. Avion subfuscus 
(young). 

13 Testacella maugei. 



PLATE 1 

The scale lines represent one centimetre. 

3. Avion atev atev. 

7. Avion intevmedius 

(contracted) . 

10. Testacella haliotidea. 



12. Milax g agates 
(contracted) . 



2. Avion subfuscus. 

5. Avion fasciatus. 
8. Avion fasciatus 
(contracted) . 

11. A vion atev vufus 
(young). 
14. Lehmannia poivieri. 




Bull. B.M. {N.H.) Zool. 6, 3 



PLATE 1 






1 \ 



PLATE 2 

The scale lines represent one centimetre. 
15. Agriolimax reticitlatus. 16. Limax tenellus. 

17. Agriolimax caruanae. 18. Lehmannia marginata. 

19. Agriolimax laevis. 
20. Milax budapestensis. 21. Lehmannia marginata var. tigrina. 

22. Limax flavns. 23. Milax sowerbyi. 

(contracted) . 
24. Limax maximns. 
25. Limax cinereoniger. 



Bull. B.M. (N.H.) Zoo'L 6, 3 



PLATE 2 



15 



^^~^=^-* 



-16 



19 




20 



21 




22 





1 1 




A REVISION OF THE LAKE 

VICTORIA HAPLOCHROMIS SPECIES 

(PISCES, CICHLIDAE) 

PART IV 




GREENWOOD 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY Vol. 6 No. 4 

LONDON : i960 



A REVISION OF THE LAKE VICTORIA 

HAPLOCHROMIS SPECIES (PISCES, CICHLIDAE) 

PART IV 



BY 

P. H. GREENWOOD 

Department of Zoology, British Museum (Natural History) 



Pp. 227-281 ; 21 Text-figures 




BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ZOOLOGY Vol. 6 No. 4 

LONDON: i960 



THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical series. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

This paper is Vol. 6, No. 4 of the Zoological series. 



Trustees of the British Museum, i960 



PRINTED BY ORDER OF THE TRUSTEES OF 
THE BRITISH MUSEUM 

Issued March, i960 Price Twenty Shillings 



A REVISION OF THE LAKE VICTORIA 
HAPLOCHROMIS SPECIES (PISCES, CICHLIDAE) 

PART IV 

By P. H. GREENWOOD 

Department of Zoology, British Museum (Natural History) 



CONTENTS 



Introduction .... 

Haplochromis lacrimosus (Blgr.) 

Haplochromis pallidus (Blgr.) . 

Haplochromis macrops (Blgr.) . 

Haplochromis cinereus (Blgr.) . 

Haplochromis niloticus nom. no v. 

Haplochromis martini (Blgr.) . 

Haplochromis humilior (Blgr.) . 

Haplochromis riponianus (Blgr.) 

Haplochromis saxicola sp. nov. 

Haplochromis theliodon sp. nov. 

Haplochromis empodisma sp. nov. . 

Haplochromis obtusidens Trewavas 

Haplochromis pharyngomylus Regan 

Haplochromis ishmaeli Blgr. 
Summary ..... 
Acknowledgments 
References .... 



Page 
229 
230 

233 
236 

239 
243 
245 
248 

252 
256 
260 
262 
266 
270 

275 
280 
280 
280 



INTRODUCTION 

In previous parts of this series I have revised species-groups possessing common or 
related structural peculiarities and, in most cases, similar feeding habits. The present 
paper deals with a greater variety of structural and trophic types and many of the 
species show no obvious relationship to one another. With one exception, the 
species described below fall into three groups, namely, structurally generalized 
insectivorous species, specialized mollusc-eaters and species showing various degrees 
of structural and adaptational intermediacy between the other two groups. The 
exceptional fish, Haplochromis martini is a piscivorous predator ; it is included 
simply because of its overall resemblance to one of the insectivorous species described 
here. 

Formerly, some of the generalized species reviewed in this paper were synonymized 
with one of the mollusc-eaters (H. ishmaeli) ; the others are included because of 
various resemblances to species now resurrected and redefined. One such is Haplo- 
chromis cinereus, a species previously considered to be the extant representative of 
the ancestral type from which at least part of the present flock had evolved. These 



zool. 6, 4 



10 



2 3 o A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 

views on the central evolutionary position of H. cinereus are no longer tenable since 
the " species " thought to be H. cinereus was a complex of several distinct species, 
some more generalized than the others. Haplochromis cinereus, sensu stricto is, in fact, 
an anatomically somewhat specialized derivative from an even more generalized 
form. 

The most outstanding structural character in many of the species described below 
is an increase in the strength and size of the pharyngeal bones and musculature. 
As might be expected, these changes are reflected in the diet of the species, which 
usually include Mollusca as an important element in their food. Two species, H. 
ishmaeli and H. pharyngomylus , feed almost entirely on snails and bivalves. 

Those species with the pharyngeal mill in an intermediate stage of hypertrophy 
are able to deal with small molluscs and also with the tubicolous larvae of certain 
Trichoptera, an otherwise infrequent element in the food of insectivorous Haplo- 
chromis without strengthened pharyngeals. 

By crushing their molluscan prey within the pharynx these species stand in sharp 
contradistinction to the other groups of mollusc-eating Haplochromis in Lake 
Victoria. Species in this latter group remove the snail from its shell by holding the 
foot between the jaws and then levering the soft parts free before ingestion takes 
place (Greenwood, 1956^ and 1957). 

Haplochromis lacrimosus (Blgr.) 1906 
(Text-fig. 1) 

Tilapia lacrimosa (part) Boulenger, 1906, Ann. Mag. nat. Hist. (7) 17, 450 ; Idem, 1907, Fish. 

Nile, 515 ; Idem, 1915, Cat. Afr. Fish. 3, 234, fig. 154. 
Haplochromis cinereus (part) Regan, 1922, Proc. zool. Soc. London, 166. 

Lectotype. An adult male 76 mm. standard length (B.M. [N.H.] 1906. 5. 30. 471) 
from Entebbe. 

Description, based on 36 specimens (including the lectotype and 10 paratypes) 
66-0-97-0 mm. S.L. 

Depth of body 31-8-38-7 (M = 35*5) per cent of standard length, length of head 
30-8-35-5 (M = 33-5) per cent. Dorsal head profile straight or slightly curved, 
sloping moderately steeply. Preorbital depth 13-6-18-0 (M = 15-5) per cent of 
head length, showing weak positive allometry with standard length ; least inter- 
orbital width 20-8-26-9 (M = 23-5), snout length 26-6-32-2 (M = 29-6) per cent. 
Diameter of eye 26-1-32-6 (M = 30-4) per cent of head, ratio of eye diameter to 
preorbital depth 1*5-2-3 (mode 2-0) ; depth of cheek 17-6-23-5 (M = 20-8) per cent. 
Caudal peduncle 15-0-19-2 (M = 17-2) per cent of standard length, 1-2-1-8 (mode 
1-5) times as long as deep. 

Mouth horizontal or almost so ; jaws equal anteriorly, the lower 31-4-41-3 
(M = 37- 1) per cent of head and 1-2-2-0 (mode i-6) times as long as broad. Posterior 
tip of the maxilla extending to the vertical to the anterior orbital margin or slightly 
beyond, rarely not quite reaching the anterior orbital margin. 

Gill rakers variable, from moderately stout to slender ; 7-9 (mode 8), rarely 6 
on the lower part of the first gill arch, the lowermost two or three reduced. 



A REVISION OF THE LAKE VICTORIA HAPLOCHROMI S SPECIES 231 

Scales ctenoid ; lateral line with 31 (f.5), 32 (f.21) or 33 (f.8) scales ; cheek with 
2 or 3 (rarely 4) series ; 6 or 7 (rarely 5 J) between the lateral line and the dorsal 
fin origin, 7 or 8 (rarely 6) between the pectoral and pelvic fin bases. Scales of the 
pectoral region small or moderate. 

Fins. Dorsal with 24 (f.19) or 25 (f.16) rays, anal with 11 (f.19), 12 (f.16) or 13 
(f.i), comprising XV-XVI, 8-10 and III, 8-10 spinous and branched rays for the 
fins respectively. Pectoral 82-5-100-0 (M = 88-5) per cent of head. Caudal truncate 
or subtruncate. 

Teeth. The outer row in both jaws is composed of unequally bicuspid, relatively 
slender and sometimes slightly recurved teeth. Rarely, a few unicuspid teeth may 
occur anteriorly in this row ; likewise a few posterolateral teeth in the upper jaw 




Fig. 1. Haplochromis lacrimosus ; lectotype (from Boulenger, Fishes of the Nile). 



may be unicuspid. There are 40-60 (mode 54, modal range 48-54) teeth in the 
upper jaw. 

Teeth in the inner series are tricuspid and implanted at a slight, posteriorly 
directed angle (cf. H. cinereus where the inner teeth lie almost horizontally) ; there 
are 2 or 3 (rarely 4) rows of inner teeth in the upper jaw and 2 (less commonly 3, 
rarely 4) in the lower. 

Lower pharyngeal bone triangular and slender ; a few slightly enlarged but bicuspid 
teeth may occur in the median tooth-rows. 

Coloration. The colours of live fishes are unknown. 

Preserved material : Sexually active males. Ground colour yellowish-silver, chest 
dusky ; a dark lachrymal stripe is always present and in some specimens it may 
extend obliquely upwards through the eye and on to the nape ; there are usually 
two transverse bands across the snout. 

Two common patterns of body markings are known, (i) A large mid-lateral blotch 



232 A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 

situated slightly posterior to the pelvic fin insertion and a mid-lateral stripe running 
from a point above the second anal ray on to the caudal fin ; a faint transverse bar 
is visible immediately posterior to the edge of the operculum and two others lie 
between the mid-lateral blotch and the origin of the posterior stripe. The blotch itself 
appears to be the intensified mid-portion of a vertical bar. 

(ii) Nine, close-set and ventrally ill-defined transverse bars on the flanks ; ventrally, 
the bars tend to run into one another so that the lower region of the flank is steely- 
grey. 

All fins, except the pelvics, hyaline, the upper part of the caudal sometimes weakly 
maculate, the mid-part dark ; anal with two or three ocelli ; pelvic fins black. 

Females and quiescent males. Ground colour greyish-silver, brownish above. 
Seven to nine faint transverse bars on the flanks, not reaching the ventral or dorsal 
outlines of the body. All fins hyaline. 

Distribution. At present, H. lacrimosus is known with certainty from Lake 
Victoria ; Pappenheim & Boulenger (1914) recorded a specimen from Lake Edward, 
but I have not been able to examine their material. 



Ecology : Habitat. No precise details are available for fishes already in the col- 
lections of the B.M. (Nat. Hist.) ; specimens collected by E.A.F.R.O. come from 
only two localities, both exposed, sandy beaches with the water depth less than 
20 feet. Thus, it is impossible to generalize on the habitat preferences of H. lacri- 
mosus. 

Food. The stomach and intestinal contents of twenty fishes were examined ; 
with one exception (a fish from Entebbe) these specimens were caught at one time 
and at a single locality (Majita, Tanganyika Territory). The gut contents of the 
sixteen specimens containing food were varied. Twelve fishes contained fine sand- 
grains, bottom detritus (including fragments of plant epidermis and diatom frustules) 
and some Cladocera ; five contained remains of insect larvae (probably Diptera), 
one an adult dipt er an, one the remains of a larval Povilla adusta Navas (Ephemerop- 
tera) and one an insect egg-mass. Two fishes yielded, besides insect fragments, the 
remains of an oligochaet worm, whilst two others each contained the foot and soft 
parts of a snail. From these scanty and topographically restricted data, H. lacrimosus 
should perhaps be classified as a bottom-feeding omnivore. 

Breeding. The breeding habits are unknown. Two of the smallest fishes (male 
and female, both 66-o mm. S.L.) are adult. It seems possible that adult males reach 
a larger maximum size than do females. 

Affinities. Haplochromis lacrimosus is one of the structurally and ecologically 
generalized species of Lake Victoria. Its most striking and apparently diagnostic 
character is the markings of male fishes ; but, it must be stressed that coloration is 
known only from preserved material. In general appearance H. lacrimosus resembles 
H. pallidus (see p. 233) but the two species differ in several characters besides male 
coloration. 



A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 

Study material and distribution records 



233 



Museum 


and Reg. No. 


Locality 

Uganda 


Collector 


B.M. (N.H.).- 


—1906. 5. 30. 471 


Entebbe 


Degen 


(Lectotype Tilapia lacrimosa) 






B.M. (N.H.).- 


—1906.5.30.472-478 


,, 


,, 


,, 


1906.5.30.483-484 


Bunjako 


> »» 


,, 


1906 . 5 . 30 . 488-489 


Buganga 


,, 


fJ 


1907. 5. 7. 81-82 


Buddu Coast 


Simon 




1908. 10. 19.2-5 


Sesse Islands 


Bayon 


" 


1959.4.28.24 


Entebbe Harbour 
Tanganyika 


E.A.F.R.O 


,, 


1959.4.28. 1-23 


Majita 


• >> 



Haplochromis pallidus (Blgr.) 1911 
(Text-figs. 2 and 3) 



Tilapia pallida (part) Boulenger, 191 1, Ann. Mus. Genova (3), 5, 74 ; Idem, 1915, Cat. Afr. 

Fish. 3, 231-2. 
Labrochromis pallidus Regan, 1920, Ann. Mag. nat. Hist. (9), 5, 45 (footnote). 
Haplochromis cinereus (part), Regan, 1922, Proc. zool. Soc. London, 166. 

This synonymy is tentative, as I have been unable to locate three specimens of 
T. pallida which Regan (1922) referred to Haplochromis guiarti (Pellegrin). Regan's 
genus Labrochromis, based on a skeleton wrongly identified as T. pallida, is discussed 
on page 275. 

Description, based on twenty specimens (including the holotype and four para- 
types) 43-74 mm. S.L. 

Depth of body 33-3-38-8 (M = 35*4) per cent of standard length, length of head 
3 2 '3 _ 35'3 (M = 34- 1 ) per cent. Dorsal head profile straight or slightly curved, 
sloping at about 30°-40°. Preorbital depth 13-2-18-2 (M = 16-5) per cent of head ; 
least interorbital width 21-0-30-0 (M = 24-7) per cent. Snout as long as broad or 
slightly longer ; its length 29-1-33-4 (M = 31-2) per cent of head ; diameter of 
eye 26-1-33-4 (M = 29-5) per cent, ratio of eye diameter to preorbital depth 1-5- 
2-3 (mode 1-7) ; depth of cheek 19-0-25-0 (M = 21-7) per cent of head. Caudal 
peduncle 14-2-18-5 (M = 16-2) per cent of standard length, 1-2-1-7 (mode 1-2) times 
as long as deep. 

Mouth horizontal ; jaws equal anteriorly, the lower 32-2-40-9 (M = 37-6) per cent 
of head and 1-4-1-8 (mode i-6) times as long as broad. Posterior tip of the maxilla 
extending to the vertical through the anterior orbital margin or slightly beyond. 

Gill rakers short and stout (slender in one specimen), 7-9 (mode 9), rarely 10, on 
the lower part of the first arch, the lowermost 1-4 (or even 5) rakers greatly reduced. 

Scales ctenoid ; lateral line with 31 (f.6), 32 (f.13) or 33 (f.i) scales ; cheek with 
2 or 3 (rarely 4) series ; 6 or 7 (occasionally 5) between the lateral line and the 
dorsal fin origin, 6 or 7 (occasionally 8) between the pectoral and pelvic fin bases. 
Scales on the chest rather small. 



234 A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 

Fins. Dorsal with 24 (f.5) or 25 (f.15) rays, anal with n (f.3) or 12 (f.17), com- 
prising XV-XVI, 8-10 and III, 8 or 9 spinous and branched rays for the fins respec- 
tively. Caudal truncate ; pectoral fin 78-0-87-0 (M = 81 -o) per cent of head. 

Teeth. The outer row in both jaws is composed of unequally bicuspid, moderately 
stout teeth implanted erectly. In most fishes more than 67 mm. S.L. some unicuspid 
teeth occur postero-laterally in the upper jaw. The holotype (the largest specimen 
examined) has only weakly bicuspid teeth in the outer row. Three small specimens 
from near the Ripon Falls have somewhat more slender outer teeth than other 
specimens. There are 36-48 teeth in the upper outer series ; no clear-cut mode can 
be determined from the sample studied. 

The small and tricuspid inner teeth are implanted at a very slight angle and 
arranged in 3 (rarely 2) rows in the upper jaw and 2 or 3 rows in the lower. 




Fig. 2. Haplochromis pallidus ; holotype (from Boulenger, Ann. Mus. Genova). 



Lower pharyngeal bone triangular, usually slender but slightly thickened in three 
specimens. Most fishes have the two median rows of teeth slightly enlarged 
(especially the most posterior one or two pairs) ; in a few, all the pharyngeal teeth 
are slender. 

Coloration. The colours of live fishes are unknown. 

Preserved material. Adult males. Ground colour greyish, with faint traces of up to 
seven dark transverse bars on the flank and caudal peduncle ; branchiostegal 
membrane greyish. A distinct, vertical lachrymal stripe, continued at an angle, runs 
through the centre of the eye ; a very faint stripe runs from the posterior orbital 
margin to the angle of the preoperculum. Dorsal fin greyish, with dark lappets 
and a dark band along the basal two-thirds of the fin anteriorly, narrowing to the 
basal third or quarter posteriorly. Caudal dark proximally and along the mid-line. 
Anal hyaline, with two large, dead-white ocelli. Pelvics black, darkest laterally. 

Females and immature males. Ground colour greyish-silver, six or seven faint 
transverse bars on the flanks and, in some, two bars on the caudal peduncle ; a 
very faint lachrymal stripe. All fins hyaline. 



REVISION OF THE LAKE VICTORIA H APLOCHROMJ S SPECIES 235 

Boulenger's (191 1) account of preserved coloration differs somewhat from that 
given above, but as his material represented at least two and possibly three species, 
the discrepancies are not considered significant. 

Distribution. Known only from Lake Victoria, unless the locality " Jinja, Ripon 
Falls " for specimens nos. 1911.3.3. 127-130 implies that these fishes were caught 
below the falls in the Victoria Nile. 

Ecology. Habitat. The only precise bionomic data available are for those specimens 
collected by E.A.F.R.O. All these were from one locality, an exposed, shallow and 
sandy beach near Entebbe Airport. 

Food. Thirteen of the sixteen fishes examined had ingested matter in the stomach 
and intestine ; all these specimens were collected at one locality and at the same 




Fig. 3. Haplochromispalliclus, 
lower pharyngeal bone, oc- 
clusal view. 




02 cm 



Fig. 4. Haplochromis ripon- 
ianus ; lower pharyngeal 




bone, occlusal view. 



Fig. 5. Haplochromis saxi- 
cola ; lower pharyngeal 
bone, occlusal view. 



time. Except for three fishes, all contained moderately large quantities of bottom 
debris (sand grains, diatom frustules and fragments of plant tissue) together with 
fragmentary insect larvae (? Diptera). The exceptional specimens contained only 
bottom detritus. 

Breeding. Haplochromis pallidas is a female mouth brooder ; the two smallest 
fishes examined (a male 54 mm. S.L. and a female 58 mm. S.L.) are both adult. 

Affinities. Haplochromis pallidas must be considered one of the many small and 
generalized species in Lake Victoria. Within this group it is extremely difficult to 
suggest phyletic relationships because the degree of inter-specific differentiation is so 
slight. In general appearance H. pallidas perhaps comes nearest to H. lacrimosus, 
from which species it is distinguished by a higher modal number of gill rakers 
(9 cf. 8), fewer and somewhat stouter teeth, a lower modal eye/preorbital ratio 
(i*7 cf. 2-0) and particularly, by differences in the preserved coloration of male 



236 A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 

fishes. The nature of both oral and pharyngeal dentition suggests that a " pallidas "- 
like species could have been ancestral to the adaptational grade at present represented 
by Haplochromis humilior (see p. 252). 

Study material and distribution records 

Museum and Reg. No. Locality Collector 

Uganda 

Genoa Museum (C.E. 12912) . Jinja . Bayon 

B.M. (N.H.). — iqii .3.3. 127-130 . Jinja, Ripon Falls 

1959.4.28.25-40 . Entebbe, Airport beach . E.A.F.R.O. 

Haplochromis macrops (Blgr.) 1911 
(Text-fig. 6) 

Haplochromis stanleyi (part) Boulenger, 1914, Cat. Afr. Fish. 3, 295. 

Haplochromis ishmaeli (part) Boulenger, 1914, torn. cit. 293. 

Tilapia macrops Boulenger, 191 1, Ann. Mus. Genova (3), 5, 73. pi. Ill, fig. 1 ; Idem, 1914, Cat. 

Afr. Fish. 3, 238. 
Haplochromis macrops (part, i.e. the species as described but excluding the tentative synonymy 

of A statotilapia jeanneli Pellegrin), Regan, 1922, Proc. zool. Soc. London, 166. 

Description, based on forty specimens from Lake Victoria (including the holotype 
[Genoa Museum] and one of the paratypes) 66-91 mm. S.L. 

Depth of body 32-5-38-2 (M = 35-8) per cent of standard length, length of head 
31-0-35-1 (M = 33-0) per cent. Dorsal head profile straight or slightly curved, 
sloping at a moderate angle (ca. 35°-40°). Preorbital depth 11-5-16-3 (M = 14-2) 
per cent of head, least width of interorbital 26-6-32*2 (M = 29-7) per cent. Snout 
as broad as long or slightly broader, its length 26-6-31-0 (M = 29-0) per cent of 
head, diameter of eye 28-6-35-4 (M = 33-0) per cent, ratio of eye diameter to pre- 
orbital depth 2-0-2-9 (mode 2-3) ; depth of cheek 17*8-24-2 (M = 21-1) per cent of 
head. Caudal peduncle 14-1-18-4 (M = 16-8) per cent of standard length, 1-2-1-6 
(mode 1-4) times as long as deep. 

Mouth slightly oblique, posterior tip of the maxilla extending to the vertical to 
the anterior orbital margin or even as far as the pupil. Lower jaw 38-0-42-5 (M = 
39-5) per cent of head, 1-4-2-2 (modal range 1-7-1-8) times as long as broad. 

Gill rakers slender or, occasionally, relatively stout ; 8-1 1 (mode 9) on the lower 
part of the first arch, the lower 1-4 rakers reduced. 

Scales ctenoid; lateral line with 30 (f.6), 31 (f.17), 32 (f.13) or 33 (f.3) scales; 
cheek with 2 or 3 series. Five or 6 (occasionally 7) scales between the lateral line 
and the dorsal fin origin ; 6 or 7 (occasionally 5, rarely 8) between the pectoral and 
pelvic fin bases. Scales on the pectoral region (relative to those on the ventral 
abdominal region) moderate to large. 

Fins. Dorsal with 24 (f.6), 25 (f.30) or 26 (f.4) rays, anal with 11 (f.9), 12 (f.30) 
or 13 (f.i), comprising XV-XVII, 8-10 and III, 8-10 spinous and branched rays for 
the fins respectively. Pectoral 68-0-96-0 (M = 84-0) per cent of head. Caudal 
truncate. 



A REVISION OF THE LAKE VICTORIA HAPLOCHROMTS SPECIES 237 

Teeth. The outer row of teeth in both jaws (except those situated postero-laterally 
in the upper) is composed mainly of bicuspid, moderately stout teeth. In certain 
fishes over 80 mm. S.L. some weakly cuspidate teeth may occur. H. macrops is, 
however, unusual in that the postero-lateral teeth in the upper jaw are generally 
tricuspid in fishes less than 85 mm. S.L. ; in larger individuals these teeth may be 
unicuspid. One aberrant individual has only tricuspid teeth in the upper, outer 
row ; the corresponding row in the lower jaw has a mixture of bi- and tricuspids, 
with the former predominating. There are 46-66 (mode 60, modal range 56-60) 
teeth in the upper, outer row. 

The inner tooth-rows are made up of tri- and unicuspid teeth arranged in 2 or 3 
(rarely 4) rows in the upper jaw and 1 or 2 (less commonly 3) rows in the lower. 
Inner teeth are implanted so as to stand erect or with a very slight, posteriorly 
directed slope. 




Fig. 6. Haplochromis macrops ; holotype (from Boulenger, Ann. Mus. Genova), 



Lower pharyngeal bone triangular, not enlarged ; teeth fine and cuspidate. 

Coloration in life. Sexually active males. Ground colour dusky to intensely black. 
Dorsal fin black, lappets and margin of the soft part red, as are the spots and blotches 
between the branched rays. Anal dusky with a diffuse red flush becoming more 
intense distally : ocelli yellow. Caudal dusky, ventral half with a red flush. Pelvic 
fins dusky yellow. Quiescent males have a female-type coloration, but with yellow 
anal ocelli and red spots on the soft dorsal. Females. Ground colour greyish-yellow 
to silver-grey. Dorsal and anal fins light yellow, caudal yellowish. 

Colour of preserved material. Adult males. Dark blackish-brown, somewhat lighter, 
except in sexually active fishes, on the chest and branchiostegal membrane ; faint 
indications of six transverse bars on the flanks (generally not reaching the dorsal 
and ventral outlines) ; a dark but faint lachrymal stripe is visible in some specimens, 



238 A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 

as is a faint dark bar along the preoperculum. Dorsal fin dark (especially along the 
basal third), lappets black. Caudal fin dark. Anal dark on the basal third to half, 
pale distally, with two or three ocelli. Pelvic fins black. Females. Brownish-yellow, 
some with eight or nine faint transverse bars on the flank and caudal peduncle. All 
fins hyaline but the caudal somewhat darker. 

Distribution. Definite records of Haplochromis macrops are available only from 
Lake Victoria although there is an indication that the species may also occur in 
the Lake Edward basin. I have examined material identified as H. macrops from 
Lake Edward (one specimen B.M. (N.H.) Reg. No. 1933.2.23.397 ; see Trewavas, 
1933) and from rivers flowing into Lake Edward (see Poll, 1939 and Poll & Damas, 
1939). Of these latter specimens (twelve in all) only one (R. G. Mus. Congo 31095, 
det. David, 1936), from Rutshuru, compares closely with the Victoria population of 
H. macrops. I hesitate to identify the remaining Congo Museum specimens from 
Rutshuru (Reg. Nos. 64888-64899), but the single specimen from the B.M. (N.H.) 
seems most closely allied to H. lividus Greenwood of Lake Victoria. I have not been 
able to study the two specimens from Lake Edward (now in Berlin) which Pappen- 
heim & Boulenger (1914) identified as H. macrops. 

The single specimen (R. G. Mus. Congo 31095) from the Edward basin now referred 
to H. macrops differs slightly from the generality of Victoria fishes in having a 
somewhat larger eye and longer lower jaw ; it is an adult female, 73-0 mm. S.L. 
The principal morphometric characters of this fish are : 



D* 


H* 


Po. % 


Io.% Eye% Snt.% Ck. % 


Lj.% 


C.P.* 


Eye/Po. 


52-8 


3i'4 


15-2 


30-4 37-0 304 21-7 
* Per cent standard length. 
% Per cent head length. 


43'4 


15-0 


2-4 



Dorsal XV, 9 ; anal III, 9 ; pectoral 91-3 per cent of head. 

In characters of dentition and squamation this fish is similar to those from Lake 
Victoria. 

Ecology. Habitat. The species is apparently confined to the shallow, sandy 
regions of the lake. 

Food. The predominating food organisms in the stomachs and intestines of 
twenty-four fishes (mainly from one locality, but caught on different occasions) are 
sub-imaginal Ephemeroptera ; two fishes had, however, fed almost exclusively on 
winged termites (Isoptera) and colonial blue-green algae. Typical bottom debris 
and sand-grains usually found in the guts of other insectivorous Haplochromis were 
not recorded. The occurrence of sub-imaginal or adult insects and planktonic blue- 
green algae, together with the absence of bottom debris, suggests that H. macrops 
feeds at the surface or in mid-water. Since the algae were not digested and did not 
constitute a major proportion of the ingested matter, they may be taken accidentally 
as the fishes dart after insect prey. 

Breeding. Haplochromis macrops is a female mouth-brooder. The smallest sexually 
active fish is a female 73 mm. S.L., the smallest adult male is 78 mm. S.L. It 
appears that males attain a larger maximum adult size than do females. 



A REVISION OF THE LAKE VICTORIA HAPLOCHROM I S SPECIES 239 

Affinities. As with most of the structurally and trophically generalized Haplo- 
chr otitis of Lake Victoria, the detailed affinities of H. macrops are impossible to 
determine. The similarities existing between H. macrops and H. cinereus are discussed 
elsewhere (p. 242). The two species differ in several characters, particularly in their 
dentition and the larger eye/preorbital ratio of H. macrops (2-0-2-9, mode 2-3, cf. 
1-5-1-9, mode i-8 for H. cimreus). Also, the gill rakers of H. macrops are finer and 
more numerous (mode 9) than in H. cinereus (mode 7). 

The large eye and shallow preorbital of H. macrops serve to distinguish the species 
from most other members of the " generalized species " group. 



Study material and distribution records 




Museum and Reg. No. 


Locality 

Uganda 


Collector 


Genoa Museum (C.E. 12928) 


Bussu 


Bayon 


(Holotype) 






B.M. (N.H.).— 1911.3.3.137 


Bussu 


Bayon 


(Paratype) 






B.M. (N.H.).— 1911.3.3.114-115 


Jinja, Ripon Falls 


,, 


1959. 4. 28. 51-78 


Beach near Nasu Point 


E.A.F.R.O 


1959.4.28.79-84 


Buka Bay 

Tanganyika 


. 


1959.4.28.85 


Mwanza, Capri Bay 


,y 


1959.4.28.86 


Majita 


,, 


1959.4.28.87 


Beach near Majita 


• 



Haplochromis cinereus (Blgr.) 1906 
(Text-fig. 7) 

Paratilapia cinerea Boulenger, 1906, Ann. Mag. nat. Hist. (7), 17, 439 ; Idem, 1907, Fish. Nile, 

478 ; Idem, 1915, Cat. Afr. Fish. 3, 344. 
Haplochromis stanleyi (part), Boulenger, 1915, torn, cit., 295. 
Tilapia lacrimosa (part) Boulenger, 1906, Ann. Mag. nat. Hist. (7), 17, 450 ; Idem, 1915, torn. 

cit., 234. 
Haplochromis cinereus (part), Regan, 1922, Proc. zool. Soc. London, 166. 
Haplochromis melanopus (part), Regan, 1922, op. cit. 165. 

Description, based on twelve specimens (including the holotype) 71-81 mm. S.L. 

Depth of body 34-6-39-0 per cent of standard length, length of head 30-8-37-3 
(M = 347) per cent. Dorsal head profile straight or slightly curved, sloping at 
ca. 40°-50°. Preorbital depth 15-0-18-0 (M = 16-4) per cent of head length, least 
interorbital width 23-3-28-0 (M = 25-3) per cent. Snout as long as broad or slightly 
longer, its length 29-2-34-6 (M = 32-2) per cent of head, eye diameter 26-2-32-0 
(M = 28-7) per cent, ratio of eye diameter to preorbital depth 1-5-1-9 (mode i-8). 
Depth of cheek 20-8-26-0 (M = 23-0) per cent of head. Caudal peduncle 15-7-18-7 
per cent of standard length, 1-4-1-6 times as long as deep. 



240 A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 

Mouth horizontal or very slightly oblique ; jaws equal anteriorly, the lower 
34-6-41-3 (M = 377) per cent of head and 1-4-1-8 (mode i-6) times as long as 
broad. Lips slightly thickened. Posterior tip of the maxilla extending almost to 
the vertical through the anterior orbital margin or as far as the eye. 

Gill rakers moderately stout, 7-9 (mode 7) on the lower part of the first gill- arch, 
the lowermost two or three rakers reduced. 

Scales ctenoid ; lateral line with 30 (f.i), 31 (f.i), 32 (f.4) or 33 (f.6) scales ; cheek 
with 3 or 4 series ; 5— 6J scales between the lateral line and the dorsal fin origin ; 
7 or 8 (rarely 6 or 9) between the pectoral and pelvic fin bases. Scales on the pectoral 
region small or, less frequently, moderate. 

Fins. Dorsal with 24 (f.8) or 25 (f.4) rays, anal with n (f.4) or 12 (f.8), comprising 
XV or XVI, 9 or 10 and III; 8 or 9 spinous and branched rays for the fins respectively. 
Caudal truncate. Pectoral 72-5-92-0 (M = 80-2) per cent of head. 




Fig. 7. Haplochromis cinereus ; holotype (from Boulenger, Fishes of the Nile). 



Teeth. Teeth in the outer row of both jaws are weakly bicuspid or unicuspid, 
relatively stout and slightly recurved ; in most fishes there is an admixture of both 
types, with unicuspids predominating. There are 40-54 teeth in the upper, outer 
row. 

The inner series are composed of either unicuspid or bicuspid teeth ; less commonly 
there is a mixture of both types. A characteristic feature of the inner tooth-rows is 
the way in which the teeth are implanted obliquely so that the crowns point 
posteriorly. Three or 4 (rarely 2) inner rows occur in the upper jaw and 2 or 3 in 
the lower. 

The dentition of H. cinereus is unlike that of other species in the " generalized 
species ' ' group ; it is, indeed, typical of the dentition found throughout the large 
group of piscivorous predators. However, in other characters (syncranial architec- 
ture, feeding habits and body-form) H. cinereus is one of the generalized species 
(see p. 242). 



A REVISION OF THE LAKE VICTORIA HAPLOCHROM IS SPECIES 241 

Lower pharyngeal bone triangular, the dentigero'us surface slightly broader than 
long. Teeth in the two median rows are slightly enlarged in five of the specimens 
examined, but are slender in the remaining seven fishes ; the other pharyngeal 
teeth are slender in all specimens. 

Coloration in life unknown. Colour of preserved fishes : Adult males. Ground 
colour greyish-brown ; chest and branchiostegal membrane dusky ; a faint lachrymal 
stripe. All fins hyaline, with very faint indications of dark maculae on the soft 
dorsal ; dorsal lappets dark. Anal fin with two or three whitish ocelli (orange 
surrounded by red in newly preserved material, according to Boulenger). Pelvics 
black except for a large light area extending over the distal half but not including 
the first branched ray. Females. Brownish, silvery grey ventrally, with eight or 
nine dark transverse bars on the flanks and caudal peduncle. All fins hyaline, the 
basal third to half of the caudal weakly maculate. 

Distribution. Known only from Lake Victoria. 

Ecology. No ecological data (except the locality) are available for the three 
specimens collected by Degen. The remaining nine fishes were caught in water less 
than 20 feet deep, over a sandy bottom on both protected and exposed shores. 

Food. The stomach and intestinal contents of ten specimens (from four localities) 
were examined. Of these, three were empty and the remainder yielded sand-grains, 
bottom detritus (including fragments of plant epidermis) and some larvae of dipterous 
insects. 

Breeding. The species is a female mouth-brooder. The smallest specimens available 
(a male and a female 71 mm. S.L.) are both sexually mature. As far as can be 
determined from this inadequate sample, both sexes reach the same adult size. 

Discussion of affinities and synonymy. In Regan's revision of the Lake Victoria 
Haplochromis (1922) the definition of H. cinereus was expanded to embrace a number 
of small and generalized or near-generalized forms previously recognized as distinct 
species. The first attempt to prune this complex was Lohberger's resurrection of H 
riponianus (see p. 252). Now, with more specimens available and some knowledge 
of Haplochromis in nature, it is clear that Regan's definition of H. cinereus must be 
narrowed considerably and that a further two species (H. lacrimosus and H. pallidus) 
should be resurrected. Amongst the group of anatomically and trophically un- 
specialized Haplochromis in Lake Victoria, H. cinereus is unusual because of its oral 
dentition (see p. 240). Relatively stout, clearly bicuspid teeth in the outer series of 
the jaws and erect tricuspid inner teeth are usual in the generalized species. The 
dentition of H. cinereus, on the other hand, shows a marked tendency for slender, 
unicuspid teeth to predominate in the outer rows ; the few bicuspid teeth present 
are weakly cuspidate. The inner teeth of H. cinereus are also atypical for the group 
in that the usual erect and tricuspid form is largely replaced by slender unicuspids 
implanted so as to point posteriorly. In fact, the dentition of H. cinereus is very 
like that of many predatory species. With so few specimens of H. cinereus known 
it is impossible to generalize on its feeding habits ; however, the gut contents of 



242 A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 

seven fishes from four different localities do not even hint at the species being a 
piscivorous predator. 

At this point it should perhaps be stressed that my observations are confined to 
adult fishes 70-90 mm. long ; juveniles have still to be discovered. 

Because " Haplochromis cinereus " had become something of a dumping ground 
for any small Haplochromis species or specimen, the published information (Graham, 
1929) on distribution and habitats can no longer be considered reliable. I have 
examined " H. cinereus " material collected by Graham and find that none of these 
specimens is referable to H. cinereus, sensu stricto. The bulk of this material is of 
undescribed species and will be dealt with in subsequent papers. Thus, Graham's 
remark that " the species (H. cinereus) is therefore widely distributed except in the 
deepest part of the lake " and Brooks' (1950, p. 159) elaboration of these data do 
not apply to H. cinereus but rather to the whole species-complex of generalized, 
bottom feeding Haplochromis in the lake. 

Haplochromis cinereus has been cited as representing the ancestral type from 
which the present species-flock could have evolved (Regan, 1922 ; Greenwood, 1951). 
For the reasons mentioned above this concept must be abandoned ; there are several 
other Haplochromis species still surviving in Lake Victoria which are structurally 
closer to the ancestral type, for example H. lacrimosus, H. nubilus or H. pallidus. 

The particular affinities of H. cinereus are difficult to determine. In gross anatomy 
and appearance H. cinereus does not differ markedly from the majority of small 
Haplochromis ; only when its dentition is considered does the difference appear 
striking. Like so many members of the generalized group, H. cinereus seems to be 
an independent offshoot from one of the basic stocks. 

In overall appearance and perhaps in at least some ecological requirements 
Haplochromis macrops is the one extant species most like H. cinereus. The two 
species are differentiated principally by the larger eye, shallower preorbital and the 
stouter, more numerous outer teeth of H. macrops. 

Haplochromis cinereus is not, as Regan suggested, closely related to H. ishmaeli. 
To stress this supposed relationship, Regan noted that six of the H. ishmaeli syntypes 
were actually specimens of H. cinereus. As a result of this present revision none of 
these specimens is still retained in H. cinereus. Haplochromis ishmaeli belongs to a 
distinct phyletic line, discussed more fully on pages 269 and 273. 

Study material and distribution records 



Museum and Reg. No. 


Locality 
Uganda 




Collector 


B.M. (N.H.). — 1906.5.30.292 


Buganga 




Degen 


(Holotype) 








B.M. (N.H.).— 1906.5.30.350 


Entebbe 




>> 


1906.5.30.482 


Bunjako 




,, 


1959. 4. 28. 41 


Entebbe, Harbour 


E.A.F.R.O 


1959.4.28.48-50 


Entebbe, Airport 
Tanganyika 


beach 


" 


1959.4.28.42-47 


Mwanza, Capri 


Bay 


E.A.F.R.O 



A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 243 

Haplochromis niloticus nom. nov. 
(Text-fig. 8) 

Tilapia bayoni Boulenger, 191 1, Ann. Mus. Genova (3), 5; 72, pi. Ill, fig. 2 {nee Paratilapia 
bayoni [= Haplochromis bayoni (Blgr.), see Regan, 1922, p. 176] Blgr., 1909, Ann. Mus. 
Genova (3), 4, 304, fig. 

Haplochromis humilior (part), Regan, 1922, Proc. zool. Soc. London, 169. 

Description, based on the holotype (100 mm. S.L.) and two other specimens 96 and 
102 mm. S.L. 




Fig. 8. Haplochromis niloticus ; holotype (from Boulenger, Ann. Mus. Genova). 



The principal morphometric characters are given below : 



S.L. 


Depth* 


Head* 


Po. % 


Io. % 


Snt. % 


Eye % 


Ck. % 


Lj. % 


C.P.* 


96-0 


35'4 


33-3 


17-2 


21-8 


31-3 


3i-3 


20-4 


39-o 


18-2 


fiooo 


33-o 


33-o 


15-2 


24-3 


3o-3 


33'5 


24-0 


33-3 


17-5 


I02-0 


34*3 


33-8 


17-4 


24-7 


29-0 


30'5 


23-2 


37*7 


17-1 



f Holotype. 

% Per cent, of head length. 

* Per cent, of standard length. 

Dorsal head profile sloping rather steeply (ca. 45°-50°) and slightly curved. Mouth 
horizontal ; jaws equal anteriorly, the length/breadth ratio of the lower 1-6-1-9. 
Posterior tip of the maxilla extending to the vertical through the anterior part of 
the eye. Lips not markedly thickened. 



ZOOL. 6, 4 



17 



244 A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 

Gill rakers moderately stout ; 7 or 8 (the lowermost one or two reduced) on the 
lower part of the first arch. 

Scales ctenoid ; 32 in the lateral line ; cheek with 2 or 3 series. Six or 7 scales 
between the lateral line and the dorsal fin origin, 8 or 9 between the pectoral and 
pelvic fin bases ; scales on the pectoral region small. 

Fins. Dorsal with 25 (f.2) or 26 (f.i) rays, anal with 12, comprising XVI, 9-10 
and III, 9 spinous and branched rays for the fins respectively. Pelvic fins with the 
first branched ray only slightly produced and extending to the origin of the anal. 
Caudal sub-truncate, scaled only on its proximal half. 

Teeth. The outer row in both jaws is composed of moderately slender, movably 
implanted and unequally bicuspid teeth ; the most posterior five or six teeth in 
the upper jaw are caniniform and stouter than those situated anteriorly and 
laterally. There are 65-70 teeth in the upper, outer series. 

Teeth in the inner series are small and tricuspid, and are arranged in four or five 
rows in the upper jaw and four in the lower. The interspace between the inner and 
outer series is very narrow. 

Lower pharyngeal bone triangular, slender or slightly enlarged, the two median 
rows of teeth relatively coarse in two specimens (including the holotype) and some- 
what more enlarged in the third. In the latter fish the next lateral row of teeth is 
also enlarged and the most posterior teeth of the median rows are sub-molariform. 

Osteology. A complete skeleton was prepared from one of the specimens caught at 
the same time as the three fishes described above. However, since H. niloticus is 
very similar to H. nuchisquamulatus (which also occurs in the Victoria Nile) it is 
difficult to confirm the specific identity of the skeleton. Apparently the sole diag- 
nostic osteological character is the lower pharyngeal bone, which is slender in H. 
nuchisquamulatus and slightly thickened in H. niloticus. The lower pharyngeal of 
the skeleton is that of H. niloticus and on this character alone the skeleton is referred 
to H. niloticus. In all other characters, except the oral dentition, the skeleton of 
H. niloticus resembles that of a generalized Haplochromis species. There are 14 + 16 
vertebrae. 

Coloration of live fishes is unknown. The three preserved specimens are all ap- 
parently males (judging from the well-defined anal ocelli) and adult. Because most 
of the coloration is now lost (the fishes are a uniform brownish-grey) I quote the 
description given by Boulenger (191 1) of the then newly preserved specimens. 
" Back dark olive to blackish, sides brassy yellow to coppery red ; a more or less 
distinct black bar below the eye ; dorsal and ventrals brown to black ; anal pink, 
blackish at the base, usually with two or three large orange ocellar spots ; caudal 
brown or blackish the lower third often pink." From this description the coloration 
of male fishes seems to be remarkably like that of H. humilior from Lake Victoria 
(see p. 250). 

Distribution. Haplochromis niloticus is known only from the Victoria Nile ; no 
information is available on its habitat or on feeding and breeding habits. 

Affinities. Haplochromis niloticus has been compared with two Haplochromis 
species from Lake Victoria. In his original description, Boulenger compared the 



A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 245 

species with H. martini, whilst Regan (1922) considered H. niloticus to be conspecific 
with H . humilior. In my opinion H. martini and H. humilior are not closely related 
to one another and H. niloticus is not allied to either. The three species differ in 
several fundamental characters, especially in the nature of their dentition. Ad- 
mittedly, the somewhat enlarged median pharyngeal teeth of H. niloticus approach 
the condition found in some specimens of H. humilior, but the oral dentition of 
the two species is very dissimilar. The outer teeth are finer and more numerous in 
H. niloticus and there are more rows of inner teeth barely separated from the outer 
row. These same characters, together with a somewhat different arrangement of 
the jaw skeleton, serve to separate H. niloticus from H. martini. 

The nature of the dentition in H. niloticus suggests affinity with H. nigricans and 
H. nuchisquamulatus , particularly the latter. From H. nigricans, H. niloticus is 
distinguished by its more generalized neurocranium and dentary (see Greenwood, 
19566) and its slightly enlarged lower pharyngeal teeth. From H. nuchisquamulatus, 
H. niloticus is again distinguished by having somewhat enlarged pharyngeal teeth 
and by a narrower interorbital region. 

The affinities of H. niloticus are not especially obvious ; the species is probably 
yet another slightly specialized side branch from the generalized Haplochromis stem. 

Study material and distribution records 



Museum and Reg. No. 


Locality 

Uganda 


• 


Collector 


Genoa Museum (C.E. 12932) 


Kakindu, Victoria Nile 


. 


Bayon 


(Holotype T. bayoni Blgr. 191 1) 








B.M. (N.H.).— 191 1.3 -3- 124-5 


tt tt 


. 


tt 


1911.3.3.126 


n n 


. 


a 


(skeleton) 









Haplochromis martini (Blgr.) 1906 
(Text-fig. 9) 

Tilapia martini, Boulenger, 1906, Ann. Mag. nat. Hist. (7), 17, 449. 
T. martini (part), Idem, 1914, Cat. Afr. Fish. 3, 239, fig. 158. 
Haplochromis martini, Regan, 1922, Proc. zool. Soc. London, 171. 

Regan (1922) based his redescription of H. martini on three of the six syntypes, 
but did not indicate to what species he referred the remaining type specimens. 
However, amongst the material Regan identified as H. cinereus there are three 
type specimens of T. martini (B.M. (N.H.) Reg. No. 1906.5.30.466-468). Presum- 
ably it was Regan's intention to include these in the published synonymy of H. 
cinereus. The three specimens are not H. cinereus but are, in fact, Haplochromis 
martini. 

Lectotype. A brooding female 88 mm. standard length (B.M. (N.H.) Reg. No. 
1906.5.30.465) from Bunjako, Uganda. 

Description, based on twenty-nine specimens (including the lectotype and four 
paratypes) 59-104 mm. S.L. 



246 A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 

Depth of body 30-8-38-0 (M = 34-4) per cent of standard length, length of head 
31-0-38-1 (M = 35-4) per cent. Dorsal head profile very strongly decurved, the 
snout sloping at an angle of ca. ^o°-yo°. Depth of preorbital 13-0-20-0 (M = 16-6) 
per cent of head, least interorbital width 20-4-26-8 (M = 24-1) per cent. Snout 
slightly longer than broad, its length 27-3-34-4 (M = 30-2) per cent of head ; dia- 
meter of eye 29-4-37-5 (M = 317) per cent, depth of cheek 20-4-27-7 (M = 24-6) 
per cent. Caudal peduncle 15-3-20-6 (M = 17-3) per cent of standard length, 1-2-1-8 
(mode i-6) times as long as deep. 

Jaws equal anteriorly, the lower 38-4-45-8 (M = 42-6) per cent of head, I-6-2-I 
(modal range 1-7-2-0) times as long as broad ; mouth horizontal or slightly oblique. 
Posterior tip of the maxilla extending to the vertical through the pupil or, less 
commonly, only to the anterior part of the eye. Such a marked posterior extension 
of the maxilla is unusual in Lake Victoria Haplochromis and may be considered 
one of the diagnostic characters of H. martini. 




Fig. 9. Haplochromis martini ; lectotype (from Boulenger, Fishes of the Nile). 



Gill rakers on the first arch moderately stout (but rather slender in a few specimens), 
one or two of the uppermost often flattened and bifid ; 8 or 9, rarely 7 or 10 
(mode 9) gill rakers on the lower part of the arch. 

Scales ctenoid; lateral line with 31 (f.5), 32 (f.7), 33 (f.9), 34 (f.7) or 35 (f.i) 
scales ; cheek with 3 or 4 rows. Six to 8 scales between the lateral line and the 
dorsal fin origin, 7 or 8 (rarely 6) between the pectoral and pelvic fin bases. 

Fins. Dorsal with 24 (f.9), 25 (f.19) or 26 (f.i) rays, anal with n (f.8), 12 (f.19) 
or 13 (f.2), comprising XIV-XVI, 8-10 and III, 8-10 spinous and branched rays 
for the fins respectively. Pectoral fin 73*5-86-5 (M = 82-0) per cent of head ; caudal 
truncate or sub-truncate. 

Teeth. In the outer row of both jaws the teeth are slender and mainly unequally 
bicuspid in fishes less than 85 mm. S.L. Above this size both weakly bicuspid and 
unicuspid teeth occur together. In fishes of all sizes a few tricuspid teeth are found 
postero-laterally in the upper jaw, an uncommon character in Lake Victoria Haplo- 
chromis (see H. macrops, p. 237). There are 46-76 teeth in the upper jaw (ill-defined 
modal range 68-70). 






A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 247 

Tricuspid teeth predominate in the inner series although in one large fish (101 mm. 
S.L.) the inner teeth are all unicuspid. The inner series are arranged in 2 (less 
commonly 3) rows in the upper jaw and in 1 or 2 (less commonly 3) rows in the 
lower. 

Lower pharyngeal bone triangular, not enlarged and with slender, cuspidate teeth. 

Osteology. The neurocranium does not differ greatly from the generalized Haplo- 
chromis type, but the premaxilla has relatively longer dentigerous arms. 

Coloration in life. Sexually active males. Ground colour golden-yellow, shading to 
silvery- white, with three or four faint black blotches below the insertion of the 
dorsal fin, and a distinct coppery sheen on the nape and anterior part of the flank. 
Dorsal fin hyaline, with a pinkish flush. Distal half of the anal scarlet, the proximal 
half colourless ; anal ocelli orange-red. Caudal fin flushed with scarlet, especially 
intense on the distal half. Pelvics dark on the outer half, reddish-yellow mesially. 
Quiescent males golden-yellow shading to silvery-white ventrally ; a fairly distinct 
dark mid-lateral stripe and an interrupted upper stripe slightly below the insertion 
of the dorsal fin. Dorsal and caudal fins darkish, the anal and pelvics lighter ; anal 
ocelli yellow. Females and juvenile males. Ground colour and banding as above, 
the upper band usually broken into rather indistinct blotches. All fins light yellow, 
the caudal somewhat darker. 

Preserved material. Both sexes. Ground colour yellowish-silver to brownish, an 
intense, narrow mid-lateral black line extends from the upper angle of the oper- 
culum to the caudal peduncle and, in some specimens even on to the caudal fin ; a 
fainter, often interrupted black stripe runs mid-way between the upper lateral line 
and the dorsal fin base. In some fishes there are traces of a very faint interocular 
band. All fins hyaline ; in males the dorsal has dusky lappets and the pelvics are 
dark. 

Distribution. Haplochromis martini is known only from Lake Victoria. Specimens 
from Lake Edward once identified as Tilapia martini (Boulenger, 1914) were later 
referred to H. schubotzi (Regan, 1921). 

Ecology. Habitat. Available records (from eight localities) suggest that H. martini 
is restricted to water less than 40 feet deep, where the species is ubiquitous but 
nowhere common. The species has been found over both sand and mud bottoms, 
on exposed shores and in sheltered bays. There are some indications that it may not 
occur close inshore since the only record of H. martini in beach-operated seine nets 
came from an area (Majita) where the nets were shot about 300 yards off-shore. 

Food. Sixteen of the twenty-two specimens examined had ingested material in 
the stomach and intestines ; in three of these specimens, however, the contents were 
unidentifiable sludge. Eleven of the remaining thirteen fishes contained, as the 
exclusive or predominating food, the fragmentary remains of small fishes (identified 
in two cases as Haplochromis) ; one of these individuals had also fed on larval 
Diptera and another had eaten what appeared to be the foot and other soft parts 
of a snail. Another fish had fed only on larval Diptera and one was empty except 
for some small fish bones in the posterior intestine. 



248 A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 

Breeding. Haplochromis martini is a female mouth-brooder ; females reach sexual 
maturity at ca. 80 mm. S.L. ; no data are available for males. Both sexes appear to 
attain the same maximum adult size. 

Affinities. Because of its strongly decurved snout, large eye and the marked 
posterior extension of the premaxilla, H. martini is one of the more immediately 
recognizable species. Yet, despite these characters H. martini retains most of the 
fundamental features of a generalized species such as H. macrops. On the other hand, 
H. martini differs from members of this species group (and probably most other 
Lake Victoria species) in its bright yellow coloration. 

When attempting to assess the phyletic affinities of H. martini one is faced with 
these rather contradictory characters and with the fact that, despite its generalized 
dentition and body-form, H. martini can be a piscivorous predator. The majority 
of piscivorous Haplochromis in Lake Victoria are larger than H. martini and have 
elongate bodies and heads ; the teeth in these species are usually large and canini- 
form. 

Taking into account the various structural and ecological characters H. martini 
should perhaps be considered a superficially but trophically distinct branch from the 
generalized stem as represented, perhaps, by an H. macrops-like ancestor. 

Study material and distribution records 

Museum and Reg. No. Locality . Collector 

Uganda 

B.M. (N.H.). — 1906.5.30.463-5 . Bunjako . Degen 

(Lectotype and paratypes) 

1906.5.30.466-8 
(Paratypes) 

1959.4.28. 124-132 . Pilkington Bay . E.A.F.R.O. 

1959.4.28. 138 . Old Bukakata Bay 

1959.4.28. 140 . Napoleon Gulf, near Jinj a 

Tanganyika 
1959.4.28. 1 16-123 . Majita 

Kenya 

,, 1959.4.28. 133-137 . Off Port Southby . ,, 

„ 1959.4.28. 139 . Beach below Usoma Lighthouse . ,, 



Haplochromis humilior (Blgr.) 1911 
(Text-figs. 10 and 11) 

Tilapia humilior Boulenger, 191 1, Ann. Mus. Genova (3), 5, 74, pi. Ill, fig. 3. 
Tilapia lacrimosa (part), Boulenger, 1915, Cat. Afr. Fish. 3, 234. 
Haplochromis desfontainesii (part), Boulenger, 191 5, torn, cit., 303. 
Haplochromis nubilus (part), Regan, 1922, Proc. zool. Soc. London, 164. 
Haplochromis humilior (part), Regan, 1922, op. cit., 169. 
? Paratilapia granti (part), Boulenger, 1915, torn, cit., 342. 



A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 249 

Lectotype. A male 90 mm. standard length from Kakindu, Victoria Nile, collected 
by Bayon (now in the collections of the Museo Civico di Storia Naturale, Genoa). 

Description, based on thirty specimens 65-90 mm. S.L. (including the lectotype 
and one paratype ; the second paratype [B.M. (N.H.) Reg. No. 1911.3.3.152] is 
very poorly preserved and although examined, is not included in the description). 

Depth of body 29-0-37-5 (M = 34-4) per cent of standard length, length of head 
31-6-37-8 (M = 34-7) per cent. Dorsal head profile curved, sloping at an angle of 
45°-5°°- Preorbital depth 13-6-17-9 (M = 16-3) per cent of head, least interorbital 
width 21-0-28-6 (M = 24-2) per cent. Snout as long as broad or slightly longer, its 
length 27-0-34-8 (M = 30-9) per cent of head, diameter of eye 27-0-32-5 (M = 30-3), 
depth of cheek 18-5-23-2 (M = 21-2) per cent. Caudal peduncle 15-2-19-1 (M = 17-4) 
per cent of standard length, 1-3-1-8 (mode 1-5) times as long as deep. 




Fig. 10. Haplochromis humilior, lectotype (from Boulenger, Ann. Mus. Genova). 



Mouth horizontal, the lower jaw often slightly shorter than the upper ; length of 
lower jaw 33-4-39-6 (M = 36-6) per cent of head and 1-3-2-2 (modal range 1-5-1-8) 
times as long as broad. Posterior tip of the maxilla extending to the vertical through 
the anterior orbital margin or somewhat beyond (to below the pupil in one specimen). 

Populations of H. humilior from different localities in the lake appear to have 
characteristic f acies which make any one population more or less readily identifiable ; 
unfortunately no means of quantifying these characters could be determined. Also, 
it has so far proved impossible, through lack of material, to decide whether Lake 
Victoria H. humilior differ from those inhabiting the Victoria Nile. 

Gill rakers short and stout, 6-8 (modes 6 and 7), rarely 9 on the lower part of the 
first arch. 

Scales ctenoid ; lateral line with 30 (f.7), 31 (f.7), 32 (f.7) or 33 (f.7) scales ; cheek 
with 2 or 3 (rarely 4) series. Six or 7 scales between the lateral line and the origin 



250 A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 

of the dorsal fin, 6-8 (rarely 5 or 9) between the pectoral and pelvic fin bases. Scales 
on the pectoral region relatively small. 

Fins. Dorsal with 24 (f.6), 25 (f.21) or 26 (f.3) rays, anal with n (f.4) or 12 (f.24) 
comprising XV-XVI, 8-10 and III, 8 or 9 spinous and branched rays for the fins 
respectively. Pectoral 69-0-92-3 (M = 81 -o) per cent of head. Caudal truncate. 

Teeth. The outer teeth in both jaws are moderately stout and unequally or, less 
frequently, sub-equally bicuspid. In fishes over 70 mm. S.L. some weakly bicuspid 
or even unicuspid teeth may occur ; there are 36-52 (modal range 46-48) teeth in 
the upper, outer series. 

Inner teeth are tricuspid and arranged in 2 or 3 (rarely 4) rows in the upper jaw 
and 1 or 2 (rarely 4) in the lower. A distinct space separates the inner and outer 
tooth series. 

Lower pharyngeal bone and teeth. The lower pharyngeal bone, although relatively 
stout is less massive than the bone in a specimen of H. ishmaeli or H. pharyngomylus 




02cm \Q 



Fig. 11. Haplochromis humilior ; 
lower pharyngeal bone, occlusal 
view, from a Lake Victoria speci- 
men. 




02 cm 



Fig. 12. Haplochromis theliodon ; 
lower pharyngeal bone, occlusal 
view. 



of the same size. Teeth in the two median rows are always enlarged, with the most 
posterior one or two pairs largest. There is some variation in the degree of enlarge- 
ment of the median teeth, which may be bicuspid, conical or even molariform ; no 
clear-cut correlation could be detected between, on the one hand, tooth-size and 
form and, on the other, the sex and size of the fish. There is, however, a tendency 
for larger individuals to have the coarsest median pharyngeal teeth. Sagittal 
sections through a number of bones (from fishes 65-70 mm. S.L.) suggest that both 
time and chance may influence the nature of the dentition, since unerupted replace- 
ment teeth are always molariform irrespective of the nature of the functional teeth 
which they underlie. 

In addition to the two median rows, the next lateral row on each side may also 
contain a number of molariform teeth. 

Coloration in life. Breeding males. Ground colour dark silvery-grey with intense 
dusky blotches on the head ; branchiostegal membrane dull black. A coppery 
flush extends over the cheek, operculum and flank as far as the origin of the anal 
fin. Anal and caudal fins light red, the colour becoming more intense along the 



A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 251 

margins of both fins and the upper and lower posterior angles of the caudal ; two 
or three yellow anal ocelli. Dorsal fin dusky, with an orange-red margin to the soft 
part and red spots and bars between the rays of the posterior half of the spinous 
dorsal and over the entire soft part. Pel vies black. Quiescent males. General 
coloration as in females except that the pelvics are somewhat dusky and the un- 
paired fins have a pinkish flush ; anal ocelli are present. Females. Ground colour 
silvery-yellow. Dorsal, pelvic and anal fins pale yellow, the dorsal with red spots 
distributed as in males. 

Colour of preserved material. Males. Greyish, darker dorsally, the chest and 
branchiostegal membrane sooty ; in some specimens there are traces of seven or 
eight, narrow transverse bars on the flank and caudal peduncle. A lachrymal stripe, 
a pair of transverse stripes across the snout and, in some fishes, a broad band across 
the interorbital region are also visible, as are one or two bands on the nape. Dorsal 
fin variable, from hyaline to dusky. Anal hyaline in quiescent fishes and whitish in 
active individuals. Pelvics black (darkest in active fishes) but with a whitish overlay 
on the proximal half. Females silvery, some with very faint traces of vertical bars 
usually most obvious on the mid-lateral aspects of the flanks. All fins hyaline. 

Distribution. The species occurs in Lake Victoria and the Victoria Nile. 

Ecology. Habitat. In Lake Victoria the species is confined to shallow water over 
sandy beaches in both exposed and sheltered areas. No data are available for the 
riverine populations. 

Food. The gut contents of thirty-two fishes (from two localities) yielded identifi- 
able material. Of these specimens, twenty-four contained bottom debris (sand 
grains, plant fragments, diatom frustules and blue-green algae) together with 
remains of both larval and pupal insects (especially Diptera and Trichoptera, less 
frequently, Ephemeroptera) ; four fishes contained only insect remains and one only 
the diatom Melosira. Ten individuals had eaten, in addition to insects, both bivalves 
(unidentifiable) and gastropods (Melanoides sp. and Bellamy a sp.). These molluscan 
fragments were too finely divided to allow for any estimate of the number of animals 
eaten. Nevertheless it does seem, from this sample at least, that Mollusca are not a 
major element in the food of H. humilior. 

The large quantities of sand found with the remains of Trichoptera larvae in 
most fishes is of interest, particularly since the grains are small and of a remarkably 
uniform size. This suggests that the sand grains could be derived from the sand- 
grain cases made by certain caddis larvae. The moderately large pharyngeal mill 
of H. humilior may thus serve the dual purpose of crushing mollusc shells and the 
sand-grain cases of certain larval insects. 

Breeding. Nothing is known about the breeding habits of H. humilior. The smallest 
specimens available (a male and a female both 65 mm. S.L.) are adult ; the sexes 
apparently do not differ in maximum adult size attained. 

Affinities. Haplochromis humilior differs from the generality of small Haplochromis 
species in several characters, particularly in having a relatively massive lower 



252 A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 

pharyngeal bone and in having a low modal number of gill rakers (6-7). Another 
pronounced difference lies in the tendency for the lower jaw to be shorter than the 
upper. In most other characters, H. humilior resembles H. pallidus, a species which 
shows incipient hypertrophy of the lower pharyngeal bone. Haplochromis humilior 
could be a more specialized off-shoot from an H. pallidus-like stem. Unlike many 
other presumed phyletic lines within the Victoria species flock, this one is not 
continued by one or more extant species. Haplochromis humilior does not appear to 
have any close relationship to the principal group of species with enlarged pharyn- 
geals, namely H. obtusidens, H. ishmaeli and H. pharyngomylus. 

Study material and distribution records 



Museum and Reg. No. 




Locality 

Uganda 


Collector 


Genoa Museum (C.E. 12910) 




Kakindu, Victoria Nile 


Bayon 


(Lectotype) 








B.M. (N.H.).— 1911. 3-3-152-3 




> > >> 


,, 


(Paratypes) 








1906. 5. 20. 314 




Entebbe 


Degen 


1959. 4. 28. 88-1 


07 


Beach near Nasu Point 


E.A.F.R.O. 


1959. 4. 28. 108 




Near Grant Bay 


• >> 


1959.4.28. 109- 


112 


Entebbe, Harbour 
Tanganyika 


• »» 


1959.4.28. 113- 


115 • 


Majita 
Kenya 


• >» 


1909. 1 1. 15. 38 




Kisumu Bay 


. Blayney-Percival 



Haplochromis riponianus (Blgr.) 1911 
(Text-figs. 4 and 13) 



Mus. Genova (3), 5, 69, pi. II, fig. 3 ; 



Pelmatochromis riponianus (part) Boulenger, 191 1, A: 

Idem, 1915, Cat. Afr. Fish. 3, 411, fig. 280. 
Haplochromis riponianus, Lohberger, 1929, Zool. Anz. 86, 222. 
Paratilapia serranus (part), Boulenger, 191 5, torn, cit., 334. 
Paratilapia victoriana (part), Boulenger, 1915, torn, cit., 341. 
Haplochromis ishmaeli (part) Boulenger, 1906, Ann. Mag. nat. Hist. (7), 17, 446 

torn. cit. 293. 
Haplochromis cinereus (part), Regan, 1922, Proc. zool. Soc. London, 166. 



Idem, 1915, 



On the basis of specimens in the Vienna Museum, Lohberger (1929) decided that 
Regan's views on the conspecificity of Pelmatochromis riponianus and H. cinereus 
could not be substantiated ; consequently he resurrected the former species as 
Haplochromis riponianus. I have not examined Lohberger 's specimens but, from 
studying considerably more material than was available to either Regan or Lohberger, 
I can fully endorse the latter 's action. 

Lectotype. A male 95-5 mm. standard length, from Jinja, Uganda, collected by 
Bayon (Genoa Museum, C.E. 12996). 



A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 253 

Description, based on twenty-eight specimens (including the lectotype and two 
paratypes) 57-104 mm. S.L. One other paratype (B.M. (N.H.) Reg. No. 1911. 
3 • 3 • 37) is n °t included in the description. 

Depth of body 33*3-39/4 (M = 357) per cent of standard length, length of head 
32-8-37-7 (M = 35-7) per cent. Dorsal head profile straight or very slightly curved, 
sloping at an angle of ca. 35°-45°. Preorbital depth 16-3-19-5 (M = 17-6) per cent 
of head, least interorbital width 23-0-28-1 (M = 25-1) per cent. Snout slightly 
longer than broad, its length 30-5-35-4 (M = 33-6) per cent of head ; diameter of 
eye 24-2-31-0 (M = 26-6), depth of cheek 19-2-25-0 (M = 22-7) per cent. Caudal 
peduncle 14-4-18-4 per cent of standard length, 1-2-1-7 times as long as deep (modal 
range 1-3-1 -5). 




Fig. 13. Haplochromis riponianus ; lectotype (from Boulenger, Ann. Mus. Genova). 



Mouth horizontal or very slightly oblique, jaws equal anteriorly, the lower 33-4- 
42-2 (M = 38-5) per cent of head and 1-3-2-0 (modal range 1-6-1-8) times as long 
as broad. Posterior tip of the maxilla extending to the vertical through the anterior 
orbital margin or almost so, occasionally to below the anterior quarter of the eye. 
Lips noticeably thickened but not produced into median lobes. 

Gill rakers on the first arch moderately stout in most fishes but slender in a few 
others, the lowermost one to three reduced and the pair nearest the epi-cerato- 
branchial angle often flattened and tri- or quadrifid ; 6-8 (mode 7) rakers on the 
lower part of the arch. 

Scales ctenoid ; lateral line with 30 (f.2), 31 (f.2), 32 (f.n), 33 (f.io) or 34 (f.2) 
scales, cheek with 3 or 4 series ; 5J-7 (rarely 5 or 8) scales between the lateral line 
and the dorsal fin origin ; 7 or 8 (rarely 6) between the pectoral and pelvic fin bases. 
Scales on the pectoral region moderate. 



254 A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 

Fins. Dorsal with 24 (f.3), 25 (f.24) or 26 (f.i) rays, anal with 11 (f.3) or 12 (f.25), 
comprising XV-XVI, 9-10 and III, 8 or 9 spinous and branched rays for the fins 
respectively. Pectoral 69-0-88-5 (M = 78-5) per cent of head. Caudal sub-truncate. 

Teeth. The outer teeth in both jaws are relatively slender and slightly to strongly 
recurved ; the basic cusp pattern is unequally bicuspid, but the crowns are often 
so worn that the teeth appear to be weakly cuspidate or even unicuspid and bluntly 
incisiform. In some fishes over 80 mm. S.L. initially unicuspid teeth occur and 
may even be the predominating form in fishes more than 100 mm. S.L. The number 
of teeth in the upper, outer rows shows slight positive allometry with standard 
length ; there are 38-62 teeth in this row. 

Unicuspid and weakly tricuspid teeth are found in the inner rows ; often both 
types of teeth occur together, especially in fishes over 80 mm. S.L. The inner teeth 
are implanted at an angle and may be buried in the thickened oral mucosa (possibly 
a preservation artefact). In the upper jaw, the inner teeth are arranged in 3 or 4 
(rarely 2) rows and in the lower in 2-4 (rarely 1) rows. 

Boulenger's description (191 1 and 1915) of the inner teeth as " minute " appears 
to stem from his being misled by the thickened oral epithelium which has hidden 
all but the tips of these teeth. 

Lower pharyngeal bone and teeth. The lower pharyngeal bone is triangular and in 
most specimens fairly stout. The relative degree to which the bone is enlarged is 
somewhat greater than that of the lower pharyngeal in H. humilior (see p. 250). 
In a few fishes, however, the bone is slender. This variation in stoutness is not 
entirely correlated with size since, although the smallest specimens have slender or 
but slightly thickened bones, some of the larger fishes do not have proportionately 
enlarged pharyngeals. 

The form of the teeth in the four median rows does show correlation with both 
the size of the individual and the stoutness of the bone. In fishes less than ca. 70 mm. 
S.L. some of the posterior teeth in the two median rows are enlarged ; in fishes 70- 
80 mm. most of the median teeth are enlarged as are some or all of the teeth in the 
row on each side of the two median rows. Next, in specimens above 80 mm. S.L. 
(except those with slender pharyngeal bones) the teeth of these four rows are larger 
still, whilst those in the two median rows tend to be molariform. Finally, in fishes 
over 98 mm. S.L. some of the more lateral teeth are also molariform. Large indi- 
viduals with slender lower pharyngeals have only slightly enlarged median teeth, 
comparable with those of the smaller (< 90 mm.) specimens described above. 

Osteology. The shape of the neurocranium departs from the generalized type 
towards the form found in the elongated, piscivorous predators (as typified by H. 
mento). In H. riponianus, the slope from the anterior tip of the vomer to the base 
of the supraoccipital crest is less steep and more nearly straight than in the generalized 
type of skull ; also, the preorbital region in H. riponianus is relatively longer than 
in, for example, the skull of H. obliquidens or H. macrops. 

Coloration. The colours of live fishes are unknown. 

Preserved material. Sexually active males. Ground colour greyish-brown, chest 
dusky ; a dark lachrymal stripe and faint traces of four interrupted transverse 
bars on the flanks. Dorsal, caudal and anal fins hyaline, the anal with two or three 



A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 255 

dead- white ocelli, usually arranged in a single row, but occasionally in two rows. 
Pelvic fins black. Immature males similar to females but with darker pelvics and 
small, distinct ocelli on the anal fin. Females silvery-grey shading to silver ventrally ; 
six to ten transverse bars of variable width (narrowest when most numerous) across 
the flanks and caudal peduncle ; sometimes, a very faint lachrymal stripe. All 
fins hyaline, the caudal occasionally maculate ; in a few specimens there are one 
or two, small, dead-white spots in the same position as the ocelli on the anal fin of 
males. 

Distribution. Lake Victoria and possibly the Victoria Nile. The latter locality is 
surmised from the data given in the original description of the species. In that 
paper (Boulenger, 191 1), the locality is given only as Jinja, Ripon Falls, but in the 
introduction Boulenger implies that the entire collection, of which H. riponianus 
formed part, was from the Victoria Nile, that is below the Ripon Falls. Later (Cat. 
Afr. Fish. 3, 1914) the type locality is given more specifically as " Ripon Falls, 
Victoria Nile." 

Ecology. Habitat. The species is apparently confined to sand or rock substrates 
in the littoral regions of the lake ; it has been caught in both exposed and sheltered 
localities. 

Food. Stomach and intestinal contents of thirty-two fishes (from one locality) 
show that at least this population of H. riponianus was mainly insectivorous, 
although some fishes had also fed on Mollusca. Insect larvae (especially Trichoptera 
and Ephemeroptera) were found in every fish. As in H. humilior (see p. 251) large 
quantities of uniformly sized sand-grains were found in the intestines of most fishes. 
Since these grains closely resemble those forming the cases of certain Caddis-fly 
larvae found intact in some individuals, it is suggested that the sand was derived, 
at least partly, from crushed cases. In addition to the main contents listed above, 
eight fishes contained relatively large quantities of crushed bivalve shells (? Corbicula 
sp.) ; a few fragments of gastropod shells were also found in these individuals. 

Breeding. No information is available on the breeding habits of H. riponianus. 
The smallest sexually mature fish is a female 84 mm. S.L. ; the smallest adult male 
is 86 mm. S.L. 

Affinities. In combination, the oral and pharyngeal dentition set H. riponianus 
apart from other Lake Victoria species. In appearance and in possessing a similar 
oral dentition, H. saxicola (see p. 257) appears to be the nearest relative of H. 
riponianus although it differs in having a relatively slender pharyngeal bone with 
only the two median tooth rows slightly enlarged. The two species, which differ 
somewhat in their ecological relationships, may represent separate adaptive lines 
derived from a common ancestor. 

Haplochromis pallidus seems structurally suited for consideration as the extant 
representative of the presumed ancestor. (See also p. 235.) 

Although the lower pharyngeal bone and its dentition are similar in H. humilior 
and H. riponianus and although both species have very similar ecological require- 



256 A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 

ments, the species show quite marked divergence in body-form (especially head 
shape) and oral dentition. What relationships there are probably lie far back in their 
phylogenetic history ; certainly the two species cannot be considered members of 
a recently evolved phyletic line. 

Boulenger's view (191 1) that H. riponianus and H. microdon are closely related 
can no longer be held ; the two species have very different phylogenies, as witnessed 
by several anatomical and osteological characters, and equally distinctive habits 
(Greenwood, 1959). Boulenger was undoubtedly misled by the supposedly small 
teeth of both species (Greenwood, op. cit., and above) whereas in fact the teeth of 
H. riponianus are not minute and are of a markedly different form from those of 
H. microdon. 



Study material and distribution records 



Museum and Reg. No. 


Locality 






Uganda 


Genoa Museum (C.E. 12996 

(Lectotype) 
B.M. (N.H.).— I9II.3.3-37-39 


Jinja, Ripon Falls 
>» >> 


" 


1911.3.3.24 
1906.5.30.280 


Entebbe 


" 


1906.5.30.394 
(Paratype H. ishmaeli) 
1929.8. 13. 1 


Bunjako 
Entebbe 


>> 


1959.4.28. 141-157 . 

1959.4.28. 158 

1959.4.28. 159 
1959 • 4 • 28 . 160-162 


Entebbe, Airport beach 
Hannington Bay 

Buka Bay 
Entebbe, Harbour 



Collector 
Bayon 

Degen 



Hoare 
E.A.F.R.O. 



Lake Victoria, Locality Unknown 

1 901 .6.24.86-87 . — 

1928.5.24. 136-138 . — 



Sir H. Johnson 
M. Graham 



Haplochromis saxicola,, sp. no v. 

(Text-figs. 5 and 14) 



Pelmatochromis riponianus (part) Boulenger, 191 1, Ann. Mus. Genova (3), 5, 69; Idem, 1915, 

Cat. Afr. Fish. 3, 411. 
Haplochromis cinereus (part) Regan, 1922, Proc. zool. Soc. London, 166. 

Holotype. A female in-o + 25-0 mm. long (B.M. (N.H.) Reg. No. 1959.4.28.249) 
from Ramafuta island, Uganda. 

Description, based on twenty-seven specimens (including the holotype of the 
species and two paratypes of Pelmatochromis riponianus) 106-123 mm. S.L. 

Depth of body 34-8-42-5 (M = 37-8) per cent of standard length, length of head 
35*3-42*5 (M = 37-8) per cent. Dorsal head profile straight or gently curved, 
sloping at ca. 30°-40°. Preorbital depth 15-2-19-0 (M = 17-6) per cent of head, 



A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 257 

least interorbital width 23-9-29-8 (M = 26-9) per cent. Snout slightly longer than 
broad or, less commonly, as long as broad, its length 34-0-41-8 (M = 37-5) per cent 
of head ; diameter of eye 20-4-26-8 (M = 24-4), depth of cheek 21-4-26-2 (M = 23-9) 
per cent. Caudal peduncle 13-1-17-7 (M = 15-4) per cent of standard length, i-o- 
i«5 (mode 1-3) times as long as deep. 

Mouth horizontal or very slightly oblique ; lips variably thickened but always 
noticeably enlarged. Lower jaw 39-7-46-5 (M = 43-0) per cent of head and 1-5- 
1-9 (mode i-8) times as long as broad. Posterior tip of the maxilla reaching or almost 
reaching the vertical through the anterior orbital margin, sometimes extending to 
below the eye. 

Gill rakers moderately stout, the uppermost two or three either slender or divided 




Fig. 14. Haplochromis saxicola ; holotype. Drawn by John Norris Wood. 



and somewhat flattened, the lowermost one to three usually reduced ; 7-9 (mode 8) 
rarely 6 on the lower part of the first arch. 

Scales ctenoid ; 31 (f. 2), 32 (f.15), 33 (f.9) or 34 (f.i) in the lateral line ; cheek 
with 3 or 4 series. Six or 7 (rarely 5) between the lateral line and the dorsal fin 
origin ; 6-8 (rarely 5 or 9) between the pectoral and pelvic fin bases. Scales on the 
pectoral region small. 

Fins. Dorsal with 24 (f.4), 25 (f.20) or 26 (f.2) rays, anal with n (f.i), 12 (f.24) 
or 13 (f.i), comprising XIV-XVI, 9 or 10 and III, 8-10 spinous and branched rays 
for the fins respectively. One specimen has XVI, 6 rays in the dorsal and another 
II, 8 in the anal. Pectoral 67-8-81-0 (M = 74-5) per cent of head. Caudal truncate 
or sub-truncate. 

Teeth. In the outer series of each jaw the teeth are slender, recurved and generally 
unicuspid, but a few weakly bicuspid teeth may occur in fishes less than 115 mm. 
S.L. 



258 A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 

The crowns of the teeth are often worn so as to assume a bluntly incisiform shape. 
There are 52-68 teeth in the outer row of the upper jaw (modal range 60-62). 

The inner tooth rows are composed of obliquely implanted and either unicuspid 
or both uni- and tricuspid teeth arranged in 2 or 3 (rarely 4 or 5) rows in each jaw. 

The dentition of H. saxicola bears a very strong resemblance to that of H. ripon- 
ianus, the major difference lying in the higher percentage of primarily unicuspid 
teeth in H. saxicola. However, it must be borne in mind that no available specimens 
of H. riponianus are as large as even the smallest H. saxicola. 

Lower pharyngeal bone and teeth. In ninety per cent of specimens examined, the 
lower pharyngeal bone is not enlarged ; in the remaining ten per cent it is somewhat 
thickened and resembles the lower pharyngeal of H. riponianus. 

The two median tooth rows are slightly enlarged in all specimens and may even 
be molariform in those individuals with enlarged lower pharyngeal bones. Very 
exceptionally, the median teeth are not noticeably larger than the more lateral ones. 

Osteology. The neurocranium of H. saxicola is very like that of H. riponianus 
(see p. 254) ; the premaxilla is, however, distinctive for its noticeably arched 
dentigerous arms which impart a characteristic peak to the antero-medial part of 
the bone. 

Coloration in life. Sexually active males. Ground colour dark grey-green, some 
scales on the flank with golden centres ; chest and branchiostegal membrane blacky 
a coppery-red flush on the operculum and flanks. Dorsal fin dark, lappets and 
maculae red ; caudal blue-grey with red posterior and ventral margins ; anal fin 
blue-grey, suffused with pink, especially the distal margins, ocelli bright yellow ; 
pelvic fins black. Quiescent males pale silvery-blue, almost grey dorsally ; some 
flank scales with golden centres, pectoral region silver. Dorsal, caudal and pelvic 
fins hyaline, the dorsal with pale red lappets and margin to the soft part, pelvics 
dusky on the lateral half, hyaline mesially. Females. Ground colour golden-grey ; 
all fins hyaline. 

Preserved material. Adult males (probably sexually active). Ground coloration 
dark grey-black, especially dark on the head, dorsal aspects of the flanks and along 
the ventral body surface. Lips light grey ; branchiostegal membrane black. Dorsal 
fin dark, with light maculae on the soft part. Anal fin black on the basal third, the 
distal two-thirds yellowish, with two grey ocelli. Caudal greyish ventrally, yellow 
around the margin. 

Females, juvenile and some apparently adult (? quiescent) males. Brownish-yellow 
ground coloration, darkest on the dorsal surface of the head ; four to seven, fairly 
narrow dark transverse bars on the flanks and caudal peduncle. Dorsal and caudal 
fins greyish, all other fins colourless, except for the black pelvics in adult males. 

Distribution. The species is known definitely from Lake Victoria but may also 
occur in the Victoria Nile. This locality is suggested by the presumed provenance 
of two P. riponianus paratypes now considered to be H. saxicola. For further dis- 
cussion, see under " Distribution " in H. riponianus, p. 255. 

Ecology. Habitat. The habitats in which H. saxicola has been caught are rather 
more varied than is common for most Lake Victoria Haplochromis. The species 



A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 259 

occurs in shallow water over exposed sand or shingle beaches, amongst dense plant 
stands near rocky shores and over shingle and small boulders in water 10-30 feet 
deep. Finally, one specimen was caught more than a mile off-shore in nets set on 
a mud bottom in water ca. 180 feet deep. The most consistent factors in all these 
habitats (except the latter) are the hard substrates of sand, rock or shingle. 

Food. Twenty-six fishes (from nine different localities) were examined ; of these, 
only twelve contained any ingested material in the stomach or intestine. Sand 
grains and small pebbles {ca. 2 mm. in diameter) were found in eleven individuals ; 
fragmentary insect larvae (Diptera, probably chironomids) formed the principal 
food in eight fishes and snails in one other. All specimens contained a few frag- 
mentary Ostracoda and one, an almost entire larva of the boring May-fly Povilla 
adust a. 

These very unsatisfactory data suggest that H. saxicola is a bottom feeder which 
preys on various invertebrates, particularly insect larvae. 

Breeding. No information was obtained on the breeding habits of H. saxicola ; 
the smallest specimen (106 mm. S.L.) is sexually mature. Both sexes appear to 
reach the same maximum adult size. 

Affinities. In general appearance, oral dentition, neurocranial morphology and in 
certain ecological characters, Haplochromis saxicola closely resembles H. riponianus. 
The most striking difference between the species is the typically slender lower 
pharyngeal bone of H. saxicola and the stouter pharyngeal bone and teeth of H. 
riponianus. The mean snout and lower jaw proportional lengths for H. saxicola are 
somewhat greater than those for H. riponianus. If, however, these characters are 
taken for both species and plotted against standard length, no marked discontinuity 
is observed ; in all morphometric characters H. saxicola could well be large H. 
riponianus. 

The differences in the lower pharyngeal bones and dentition, however, are against 
this interpretation, the more slender bone with but slightly enlarged median teeth 
characterizing the larger specimens described as H. saxicola. In no Haplochromis 
species with enlarged pharyngeals does the bone and its dentition become less 
coarse with growth. Indeed, the reverse is usual (see Greenwood, 1959 and p. 277). 



Study material and distribution records 



Museum and Reg. No. 


Locality 

Uganda 


Collector 


B.M. (N.H.).— 1959.4.28.249 


Ramafuta Isl. 


E.A.F.R.O 


(Holotype) 






iQir. 3-3-39 


Jinja, Ripon Falls 


Bayon 


(Paratype of Pelmatochromis riponianus) 




Genoa Museum. 






(A paratype of P. riponianus) 


)> )> 


Bayon 


B.M. (N.H.).— 1959.4.28.250-255 


Ramafuta Island 


E.A.F.R.O 


„ 1959.4.28.256-267 


>, y, 


,, 


1959.4.28.268-270 


Beach near Nasu Point 


,, 


1959.4. 28. 271-272 


Jinja 


,, 


1959.4.28.273 


o° 4' S., 33 14' E. 


t> 


zool. 6, 4 




18 



260 A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 

Haplochromis theliodon sp. nov. 
(Text-figs. 12 and 15) 

Holotype. A fish 95-0 + 20-0 mm. long (B.M. (N.H.) Reg. No. 1959.4.28. 163) 
from Majita, Tanganyika Territory. 

Description, based on seven specimens (including the holotype) 79-95 mm. S.L. 
The principle morphometric characters are summarized below : 

S.L. Depth* Head* Po. % Io. % Snt. % Eye % Ck. % L.j. % C.P.* 

75-o 37*3 37'3 i7'9 25-0 32-2 25-0 21-4 39-3 15-3 

84-0 35-7 36-9 i6-2 25-8 32-2 26-8 25-8 38-7 14-3 

85-0 36-6 36-6 i6-i 22-6 35-5 24-2 22-6 38-7 15-4 

86-o 38-4 33-7 17-3 24-1 33-5 24-1 24-1 38-0 15-1 

88-o 38-6 36-3 18-1 25-0 34-4 25-0 24-4 37-4 14-8 

89-0 38-2 36-5 18-5 24-6 33-8 24-6 27-0 38-4 15-2 

95-0 36-8 33-7 17-2 25-0 34-4 26-6 25-0 39-0 13-7 

* Per cent, of standard length. 
% Per cent, of head length. 



Dorsal head profile straight except for a slight concavity above the eye, sloping 
fairly steeply ; snout longer than broad. Mouth slightly oblique ; lips, especially 
the upper, thickened. Lower jaw 1-5-1-8 times as long as broad. Posterior tip of 
the maxilla almost reaching the vertical to the anterior orbital margin. 

Gill rakers variable, from stout (the commonest) to relatively slender ; 7 (f.i), 

8 (f.5) or 9 (f.i) on the lower part of the first arch. The lowermost two to four rakers 
may be reduced. 

Scales ctenoid ; lateral line with 31 (f.2), 32 (f.3) or 33 (f.2) scales ; cheek with 
3 or 4 series. Six to 9 scales between the lateral line and the dorsal fin origin, 7-9 
between the bases of the pectoral and pelvic fins. Scales on the pectoral region 
small and deeply embedded. 

Fins. Dorsal with 25 rays, anal with 12, comprising XV-XVI, 9 or 10 and III, 

9 spinous and branched rays for the fins respectively. Pectoral 64-3-72-0 (M = 69-5) 
per cent of head. Caudal truncate or sub-truncate. 

Teeth. In six of the seven specimens examined, the outer row in both jaws is 
composed of relatively stout, unequally bicuspid teeth, except for the most posterior 
pair in the upper jaw which are unicuspid. In the seventh specimen, the anterior 
and postero-lateral teeth are unicuspid and the lateral teeth unequally bicuspid. 
There are 36-46 outer teeth in the upper jaw. 

The inner teeth are small and tricuspid (with a few unicuspids in the exceptional 
specimen mentioned above) and are arranged in 3 or 4 rows in the upper jaw and 
3 or 4 (rarely 2) in the lower. Inner teeth are implanted vertically or somewhat 
obliquely. 

Lower pharyngeal bone and teeth. The lower pharyngeal bone is thickened and 
moderately stout ; that is to say, it is comparable with the lower pharyngeal in 
most H. humilior but finer than that in H. obtusidens (see p. 267). 



A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 261 

Five of the seven specimens examined have the four median tooth rows composed 
of large and molariform teeth, whilst the next lateral row of each side contains 
enlarged but clearly cuspidate teeth. In the exceptional fishes, only the median 
pair of tooth rows contains molariform elements and only one lateral row on each 
side has enlarged teeth. 

Coloration. The colours of live fishes are unknown. 

Preserved material. Quiescent male. Ground colour dusky bronze, with a very 
faint mid-lateral stripe, six interrupted transverse bars and a distinct lachrymal 
stripe. Dorsal fin dark, lappets pale ; caudal dark except for its pale distal margin ; 
anal pale (? orange) with four, small, white ocelli arranged in a single row. Pelvics 
black on the outer two-thirds, lighter medially. Females and immature males 
uniformly light brown except for the darker dorsal head surface and nape ; a dark 




Fig. 15. Haplochromis theliodon ; holotype. Drawn by Miss G. Osterritter. 

mid-lateral stripe of variable intensity is continuous in some specimens but 
interrupted in others above and slightly anterior to the anal fin ; when inter- 
rupted, the line is thickened to form a black blotch above the pectoral fin. A fainter, 
continuous dorsal stripe runs above the upper lateral line ; there are traces of seven 
to nine, variously interrupted transverse bars on the flanks but not on the caudal 
peduncle. Dorsal and anal fins hyaline, darker between the branched rays. Caudal 
maculate on the upper half, immaculate below. Pelvic fins hyaline in females but 
with the outer half dusky in males. 

Distribution. Known only from Lake Victoria. 



Ecology. Habitat. The few specimens known came from two localities, both 
exposed, shallow and sandy beaches. 

Food. Analysis of stomach and intestinal contents from all seven specimens 
indicate that H. theliodon is a bottom feeder with a varied diet including small 
fishes (cichlids, 20-25 mm. long), Gastropoda (foot and soft parts only), Lamelli- 
branchiata and insect larvae, e.g. Povilla adusta (Ephemeroptera) and Trichoptera 



262 A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 

(including the cases). Each fish had also ingested fairly large quantities of bottom 
debris. 

Breeding. Few data are available ; one female 75 mm. S.L. is sexually mature 
whilst another, 88 mm. S.L. appears to be a juvenile maturing for the first time. 

Diagnosis and affinities. The nature of its lower pharyngeal bone and dentition 
places H. theliodon on the same level of structural modification as H. humilior and 
H. riponianus. In other characters (especially general appearance and oral dentition) 
H. theliodon is unlike both the former species. It differs from H. humilior in several 
morphometric characters (straight and not curved dorsal head profile, deeper 
preorbital, longer snout and smaller eye) and from H. riponianus in the shape of 
the head and the nature of its oral dentition. Haplochromis theliodon also differs 
from both the other species in having much smaller and more deeply embedded 
pectoral scales. 

The affinities of H. theliodon are not easily determined except in so far as the 
species is clearly a little-modified derivative of the generalized stem. The small 
pectoral scales of H. theliodon are not common in generalized species still extant in 
Lake Victoria ; but, small chest scales do characterize certain of the generalized 
and fluviatile species of East Africa, some of which also show an incipient hypertrophy 
of the pharyngeal bones and dentition (Greenwood, unpublished). 

Study material and distribution records 

Museum and Reg. No. Locality Collector 

Tanganyika 

B.M. (N.H.).— 1959.4.28.163 . Majita . E.A.F.R.O. 

(Holotype) 

1959. 4. 28. 164-168 . „ . 

Uganda 
1959. 4. 28. 169 . J in j a Pier 



Haplochromis empodisma sp. nov. 
(Text-figs. 16 and 17) 

Haplochromis ishmaeli (part) Boulenger, 1906, Ann. Mag. nat. Hist. (7), 17, 446 ; Idem, 1915, 

Cat. Afr. Fish. 3, 293 ; Regan, 1922, Proc. zool. Soc. London, 169. 
Tilapia lacrimosa (part), Boulenger 1906, op. cit., 450 ; Idem, 1915, torn, cit., 234. 
Haplochromis cinereus (part) Regan, 1922, op. cit., 166. 

Note. Because two specimens of H. empodisma (collected during Graham's 1927- 
28 survey of Lake Victoria) were mistaken for syntypes of H. michaeli Trewavas, 
this species was referred to as H. michaeli in previous papers (Greenwood, 1954, 
1956^). All references to " H. michaeli" in these publications should now be 
corrected to read Haplochromis empodisma. 

Holotype. An adult male 117 + 23 mm. total length (B.M. (N.H.) Reg. No. 



A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 263 

1959.4.28. 170), caught on the bottom in 90 feet of water off the southern tip of 
Kibibi Island (o° 10' N. ; 33 io' E.), Uganda. 

Description, based on thirty-nine fishes (including the holotype) 65-117 mm. S.L. 

Depth of body 33*0-43-8 (M = 39-3) per cent of standard length, length of head 
33-3-39-4 (M = 36-7) per cent. Dorsal head profile straight or gently curved, sloping 
at an angle of 35°-40°. Preorbital depth 15-1-20-5 (M = 18-1) per cent of head ; 
least interorbital width 20-6-28-6 (M = 24-3), length of snout 27-5-37-2 (M = 32-9), 
diameter of eye 24-4-34-0 (M = 27-3) per cent. Depth of cheek shows a marked 
positive allometry with standard length, being 21-2-23-0 per cent of head in fishes 
less than 80 mm. S.L. (N — 4) and 23-5-31-4 (M = 27-9) per cent in larger specimens 
(N = 35). Caudal peduncle 14-5-20-0 per cent of standard length and 1-3-1-7 (mode 
1-4) times as long as deep. 

Mouth horizontal or slightly oblique ; lips not noticeably thickened. Posterior tip 
of the maxilla reaching the vertical through the anterior margin of the orbit or to 




Fig. 16. Haplochromis empodisma / holotype. Drawn by John Norris Wood. 



below the anterior part of the eye ; less frequently, not reaching as far as the orbit. 
Lower jaw 39-1-48-7 (M = 43-9) per cent of head, 1-5-2-3 (mode, ill-defined, 2-0) 
times as long as broad. Malformation of the lower jaw is relatively common in this 
species ; individuals so affected have the lower jaw broader than the upper (which 
closes within the lower) and are distinctly prognathous. 

Gill rakers usually slender, although relatively stout rakers also occur ; 7-10 
(modal range 8-9) on the lower part of the first gill arch, the lowermost 1-3 rakers 
reduced, occasionally the uppermost 1 or 2 somewhat flattened and bi- or trifid. 

Scales ctenoid ; lateral line with 30 (f.io), 31 (f.9), 32 (f.15) or 33 (f.4) scales. 
Cheek with 3 or 4 (occasionally 5) series. Six or 7 scales between the lateral line and 
the dorsal fin origin, 6-8 between the pectoral and pelvic fin bases. 

Fins. Dorsal with 23 (f.8), 24 (f.30) or 25 (f.i) rays, anal with 10 (f.i), n (f.22) 
or 12 (f.16), comprising XIV-XVI, 8 or 9 and III, 7-9 spinous and branched rays for 



264 A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 

the fins respectively. Pectoral fin 73*0-96-5 (M = 85-8) per cent of head. Caudal 
truncate or sub-truncate. 

Teeth. The teeth in the outer row of both jaws are slender and often gently 
recurved ; in fishes less than 95 mm. S.L. the teeth are unequally bicuspid but in 
larger individuals are weakly bicuspid or unicuspid ; it is usual to find both types 
of teeth in large fishes. There are 54-82 (modal range 70-72) teeth in the upper jaw. 

Teeth forming the inner series are small and usually tricuspid but some unicuspids 
may occur, particularly in fishes over 100 mm. S.L. ; there are 2 or 3 (rarely 4) rows 
in the upper jaw and 1-3 in the lower. The innermost row, especially in the upper 
jaw, is implanted obliquely. 

A common abnormality affecting both inner and outer teeth is for the crowns to 
be coarse, slightly swollen and darkly pigmented ; in such teeth the crown is globose. 

Lower pharyngeal bone and teeth. The lower pharyngeal is fine and rather narrow, 




02cm 



Fig. 17. Haplochromis empodisma ; 
lower pharyngeal bone, occlusal 




02 cm 



Fig. 18. Haplochromis obtusidens ; 
lower pharyngeal bone, occlusal 



the dentigerous surface having the outline of an isosceles triangle. Usually, none 
of the pharyngeal teeth is markedly enlarged, but in certain specimens of all sizes 
a few teeth in the two median rows are slightly coarser. Even when enlarged these 
teeth retain the same form as their lateral and more slender congeners. 

Osteology. The neurocranium and dentary of H. empodisma were figured and 
briefly described in an earlier paper (Greenwood, 1956a, p. 305, fig. 5 ; the species 
was then wrongly identified as H. michaeli). Both the neurocranium and the jaw 
elements are directly comparable with those of a generalized species such as H. 
macrops, except that in H. empodisma the outer teeth are more numerous. 

Coloration in life. Sexually active males. Ground colour dark turquoise on the 
flanks and dorsal body surface, silver-yellow ventrally ; snout and dorsal head 
surface dark red. Dorsal and caudal fins diffuse red ; anal black anteriorly, dark red 
posteriorly, the ocelli orange-red. Pelvic fins black. Quiescent males as for females 
(see below) but dorsal fin with red maculae between the branched rays. Ripening 
males show some reddening of the head and snout, whilst the maculae of the dorsal 



A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 265 

are more intense. Females. Ground colour yellowish-silver, darker dorsally. Dorsal 
and caudal fins dark neutral ; anal yellow ; pelvics very pale yellow. 

Preserved material. Males (sexually active), dark grey becoming black ventrally ; 
a distinct lachrymal stripe and two bars across the snout (quiescent males greyish 
brown, dusky on the belly ; very faint indications of up to five incomplete transverse 
bars on the flanks ; lachrymal stripe visible but no bars across the snout). Dorsal 
fin dark, the lappets black ; caudal dark proximally, lighter distally ; basal third 
of anal black, remainder of fin light, with two or three large white ocelli. Pelvic fins 
black. Females brownish-silver ; in a few specimens there are very faint indications 
of a lachrymal stripe and of incomplete transverse bars on the flanks. All fins 
hyaline ; in some fishes the upper half of the caudal is slightly maculate. 

Distribution. Known only from Lake Victoria. 

Ecology. Habitat. It seems that H. empodisma is generally restricted to those 
areas of the lake where the bottom is composed of organic mud. The species has 
been caught in bottom nets set at depths from 10-90 feet. 

Food. The stomach and intestinal contents of thirty -seven fishes (from numerous 
localities) show that H. empodisma is a bottom feeder. The principal food organisms 
are the larval (and less frequently pupal) stages of dipterous insects, together with 
diatoms derived from the bottom mud. Only two fishes yielded remains of Mollusca ; 
in one fish a few fragments of bivalve shell were found and in the other an operculum 
from a large gastropod. Since no other snail fragments were found in this fish, it is 
possible that the operculum was accidentally ingested. 

Breeding. It is not known whether H. empodisma spawns in the habitats described 
above or whether spawning takes place over a more solid substrate, for example 
outcrops of rock or sand. Females carrying embryos and larvae have been caught 
together with non-breeding fishes. The smallest sexually active fish is a female 
84 mm. S.L. ; the smallest adult male is 90 mm. S.L. Both sexes reach the same 
maximum adult size. 

Affinities. Structurally and in its feeding habits, H. empodisma must be considered 
a generalized species. It differs from the majority of generalized forms in Lake 
Victoria only by its larger size, greater number of teeth (possibly a correlate of the 
larger size) and in the wide range of depths at which it has been caught. The nearest 
living relative of H. empodisma is a small and as yet undescribed species which occurs 
in the same habitat but is confined to shallow water. 

Haplochromis empodisma and H. obtusidens are strikingly similar except for one 
structure, the lower • pharyngeal bone. In H. obtusidens the lower pharyngeal is 
thickened and carries a number of enlarged, crushing teeth, whereas in H . empodisma 
this bone is slender and carries numerous fine teeth. Haplochromis empodisma could 
well represent the ancestral type from which H. obtusidens was derived by an increase 
in size of the pharyngeal bones and a correlated change in the pharyngeal dentition. 
Although the inter-specific differences in the nature of the pharyngeal mill are fairly 
clear-cut, some individuals deviate from the specific mode in such a way as to indicate 
a likely transitional condition in the evolution of an " obtusidens "-like species. 



266 A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 

Since, in turn, H. obtusidens provides a structural type basic to the evolution of 
a specialized crushing pharyngeal mill (as found in H. ishmaeli and H. pharyngo- 
mylus), H. empodisma could represent the extant representative of the basal species 
in the phyletic line which culminated in H. ishmaeli and H. pharyngomylus. 

Study material and distribution records 



Mi 


iseum and Reg. No. 


Locality 
Uganda 


Collector 


B.M. (N.H.). — 1 959. 4. 28. 170 


Off Kibibi Island 


E.A.F.R.O. 




(Holotype) 






>> 


1906.5.30.472 


Entebbe 


Degen 


>> 


1906.5.30.402 
(Paratype, H. ishmaeli) 


Bunjako 


• 


" 


1906.5.30.404 
(Paratype, H. ishmaeli) 


• »> 


' 


,, 


1959.4.28. 180-189 


Pilkington Bay 


E.A.F.R.O. 


,, 


1959. 4. 28. 194-195 


Thruston Bay 




,, 


1959.4.28. 196-197 


Ekunu Bay 


»i 


,, 


1959.4.28. 198 


Jinja 


,, 


,, 


1959.4.28. 199 


Napoleon Gulf, near Jinja 


,, 


,, 


1959 .4.28. 202-203 


Kibibi Island 


,, 


1 


1959.4.28.358 


Off S. tip of Kibibi Island 
Tanganyika 


' 


" 


1959.4.28. 190-193 


Nyamakyamwa 

Kenya 


E.A.F.R.O. 


i " 


1959. 4. 28. 200-201 


Manadu Island 


• i > 


» 


1959.4.28. 171-179 


Off Port Southby 


• i > 




Lake 


Victoria, Locality Unknown 




»» 


1928.6.2.37-38 


— 


M. Graham 



Haplochromis obtusidens Trewavas 1928 
(Text-fig. 18) 

Haplochromis desfontainesi (part), Boulenger, 1915, Cat. Afr. Fish., 3, 302. 
Tilapia lacrimosa (part), Boulenger, 1915, torn, cit., 234. 
Haplochromis cinereus, (part) Regan, 1922, Proc. zool. Soc. London, 166. 
Haplochromis obtusidens Trewavas, 1928, Ann. Mag. nat. Hist. (10), 2, 95. 

Lectotype. An adult male 107-0 + 23-0 mm. total length (B.M. (N.H.) Reg. No. 
1928. 5. 30. 21). 

Description, based on forty-four specimens (including the lectotype and one 
paratype) 60-114 mm. S.L. 

Certain proportions show fairly well-marked allometry with standard length ; for 
these, ranges and means are given for each of the relevant size-groups. 

Depth of body 35*0-44-3 (M = 38-6) per cent of standard length, length of head 
32-5-38-2 (M = 35-8) per cent. Dorsal head profile gently curved or, less commonly, 
straight, sloping at an angle of 40°-5o°. Preorbital depth in fishes < 70 mm. S.L. 



A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 267 

(N = 4), 13-0-167 (M = 14-5) per cent of head; in larger fishes (N == 40) 15-1- 
20-5 (M = 17-9) ; least interorbital width 21-8-29-0 (M = 24-2) per cent ; snout 
length in fishes < 85 mm. S.L. (N = 6), 26-0-31-0 (M = 28-5) per cent, in larger 
individuals (N = 38), 29-0-36-4 (M = 33-3) ; diameter of eye in fishes < 85 mm., 
27-6-34-8 (M = 31-2) per cent and in larger fishes 24-3-30-8 (M = 27-2) ; depth of 
cheek in specimens < 85 mm., 18-6-23-8 (M = 21-5) and in larger individuals 21-2- 
30-0 (M = 26-7) per cent of head. Caudal peduncle 15-2-19-7 (M = 17-5) per cent 
of standard length, 1-2-1-8 (mode 1-5) times as long as deep. 

Jaws equal anteriorly ; mouth horizontal or slightly oblique ; lips not markedly 
thickened. Posterior tip of the maxilla extending to the vertical through the anterior 
orbital margin or to below the anterior part of the eye ; exceptionally, not reaching 
the orbit. Lower jaw 37-9-45-5 (M = 41-8) per cent of head and 1-3-2-0 (mode 1-6) 
times as long as broad. 

Gill rakers relatively slender, although short and stout rakers are found in a few 
individuals ; 7-9 (modal range 7-8), rarely 6, on the lower part of the first arch, 
the upper pair of rakers often flattened. 

Scales ctenoid ; 29 (f.i), 30 (f.7), 31 (f.2i), 32 (f.n) or 33 (f.4) in the lateral line ; 
cheek with 3 or 4 (rarely 5) series ; 5J-7 scales between the lateral line and the 
dorsal fin origin, 5-7 (rarely 9) between the pectoral and pelvic fin bases. 

Fins. Dorsal with 23 (f.13), 24 (f.25) or 25 (f.6) rays, anal with 11 (f.24), 12 (f.19) 
or 13 (f.i), comprising XIV-XVI, 8-10 and III, 8-10 spinous and branched rays for 
the fins respectively. Pectoral 73-5-103-0 (M = 86-8) per cent of head. Caudal 
truncate or sub-truncate. 

Teeth. In fishes less than 90 mm. S.L., the outer row in both jaws is composed of 
unequally bicuspid and relatively slender teeth. Larger individuals have an ad- 
mixture of either unequally bicuspid and unicuspid teeth or of weakly bi- and 
unicuspids. A few individuals have exclusively bicuspid or unicuspid teeth. There 
are 40-80 teeth (ill-defined mode at 70 ; modal range 66-70) in the outer row of 
the upper jaw. 

Teeth in the inner series are generally tricuspid and small ; in some fishes over 
100 mm. S.L. there may be either a mixture of tri- and unicuspid teeth or ; more 
rarely, only unicuspids. The innermost row of teeth (especially in the upper jaw) 
is implanted obliquely and in some individuals the whole inner series lie at an oblique 
angle. There are 2 or 3 (rarely 1 or 4) rows of inner teeth in the upper jaw and 1 
to 3 (usually 2, rarely 4) in the lower. 

Lower pharyngeal bone and teeth. Despite some individual variability, the lower 
pharyngeal bone of H. obtusidens is always obviously thickened and the two median 
rows of teeth are enlarged and molariform or sub-molariform. As might be expected, 
there is a positive correlation between size and the degree to which the pharyngeal 
bones and teeth are enlarged. In fishes over 90 mm. S.L. as many as six rows of 
teeth may be composed of molariform elements ; even in specimens with a few 
molariform rows the remaining teeth are enlarged, except in the upper corners of 
the bone. 

Osteology. In all respects except the form of the pharyngeal apophysis, the neuro- 
cranium of H. obtusidens resembles that of H. empodisma ; the pharyngeal apophysis, 



268 A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 

however, is stouter and broader, thereby foreshadowing the condition found in H. 
ishmaeli and H. pharyngomylus. In both these species the basioccipital facets are 
more expanded than in H. obtusidens. 

Coloration in life. Sexually active males. Ground colour light blue-black with, 
ventrally, a silver patch extending from the isthmus almost to the vent ; branchio- 
stegal membrane black. Dorsal fin dark with red lappets and margin to the soft 
part and red spots or dashes between the branched rays. Caudal dark with a blood- 
red margin and a diffuse red centre. Anal dark proximally, blood-red distally ; 
ocelli orange or orange-red. Pelvic fins black. Quiescent males as for females but 
lacking the red spots or flush on the dorsal fin. Females. Ground colour silver-grey. 
Dorsal fin dark with traces of a red flush. Caudal dark. Anal and pelvic fins yellowish. 

Preserved material. Adult males. Ground colour brownish, dusky ventrally (below 
the level of the lower lateral line), with faint indications of four transverse stripes 
originating from the dark area but not reaching the base of the dorsal fin ; the 
intensity of the dusky area varies considerably, from charcoal to coal-black. Dorsal 
fin hyaline to dusky, the basal region and the lappets black ; soft part of the fin 
sometimes maculate. Caudal dark on its proximal half, light (yellowish) distally. 
Anal dark on the basal half to third, light distally, with two large white ocelli. 
Pelvic fins black. Females silver-grey to yellowish-brown ; in some fishes there are 
very faint indications of four to six, incomplete transverse bars on the flanks. Dorsal 
fin dark or hyaline. Caudal dark, the upper half maculate in some individuals. All 
other fins hyaline. 

Note on jour atypical individuals. Four specimens, 80-94 mm. S.L. (B.M. (N.H.) 
Reg. Nos. 1959.4.28.236-239) from Old Bukakata Bay, Uganda are included in the 
description given above although they differ from the generality of specimens in 
certain characters. The lower pharyngeal bones in two of these fishes are somewhat 
more slender than is modal but are typical in the other two specimens. All four 
fishes have fewer upper outer teeth than is usual (40-52 cf. modal range 66-70) 
and somewhat shallower cheek than equivalent sized specimens from other areas. 
In this latter character, however, they resemble the two type specimens. Finally it 
must be mentioned that these fishes resemble one another in genera] facies rather 
more closely than they resemble the other specimens. Such obvious but undefinable 
and geographically localized facies are fairly common amongst Lake Victoria 
Haplochromis species. 

The exact status of the four specimens from Bukakata cannot be determined for 
want of more material from this locality. 

Distribution. Known only from Lake Victoria. 

Ecology. Habitat. Haplochromis obtusidens is predominantly a species of shallow 
water (less than 30 feet), apparently restricted to a substrate of soft, organic mud ; 
a few individuals have, however, been taken over sand and in water about 60 feet 
deep. 

Food. The gut contents of forty-six fishes (from numerous localities) indicate 
that the principal food organisms of H. obtusidens are insects (especially larval 
Diptera) and molluscs (particularly the bivalve Corbicula, although some gastropods 



A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 269 

[Melanoides] are also eaten). Together with these organisms, the fishes had ingested 
fairly substantial quantities of bottom mud, which, in the areas inhabited by H. 
obtusidens, is almost entirely composed of living and moribund diatoms. A com- 
parison of stomach and intestinal contents shows that the protoplasm of these 
diatoms is digested by the fish. Other plants, in this case mostly blue-green algae, 
are apparently undigested. 

Breeding. Haplochromis obtusidens is a female mouth-brooder ; exact spawning 
sites were not discovered. A male 83 mm. S.L. is the smallest mature fish in the 
sample studied ; the smallest mature female is 89 mm. S.L. Both sexes reach the 
same maximum adult size. 

Affinities. The relationships of H. obtusidens seem to lie with H. empodisma which 
it resembles in body-form and most syncranial characters. Haplochromis obtusidens 
differs in having an enlarged neurocranial apophysis for the upper pharyngeal bones 
and in the correlated character, enlarged pharyngeal bones and teeth. The species 
also shows certain fairly marked affinities with H. ishmaeli and H. pharyngomylus 
and could well represent the extant version of an ancestral type from which these 
two species evolved (see also p. 265). 

In Lake Edward, H. malacophagus Poll has reached a comparable evolutionary 
stage leading towards extreme pharyngeal hypertrophy, but the two species are not 
closely related. 



Study material and distribution records 




Museum and Reg. No. 


Locality 






Uganda 


B.M. (N.H.).- 


—1906. 5. 30. 531 


Buganga 


,, 


I 959-4- 28. 204-212 




Pilkington Bay 


>> 


1959. 4. 28. 213-218 




Ekunu Bay 


,, 


1959 .4.28. 228-229 




Pilkington Bay 


,, 


1959.4.28.236-239 




Old Bukakata Bay 


,, 


1959.4.28.243-244 




Dagusi Island 


,, 


1959.4.28.245 




Buka Bay 


,, 


1959.4.28.247 




Sesse Islands 



Collector 

Degen 
E.A.F.R.O. 



1909.7.27.43 

1959-4-28.219-227 

1959.4.28.230-232 

1959. 4. 28. 240-241 

1959.4.28.242 

1959.4.28.246 

1959-4-28.248 



Kenya 

Kavirondo Bay 

Off Port Southby 

Kach Bay (Kavirondo Gulf) 

Beach below Usoma lighthouse 

Off Port Southby 

Kavirondo Gulf 

Off mouth of Nzoia River 



Alluaud 
E.A.F.R.O. 



Tanganyika 

1959-4-28.233-235 . Beach near Majita 

Lake Victoria, Locality Unknown 

1928. 5. 30. 21 . — 

(Lectotype) 

B.M. (N.H.).— 1928.5.20.20 . — 

(Paratype) 



M. Graham 



270 A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 

Haplochromis pharyngomylus Regan 1929 
(Text-figs. 19 and 20) 

Haplochromis ishmaeli (part) Boulenger, 1906, Ann. Mag. nat. Hist. (7), 17, 446; 
Idem, 1915, Cat. Afr. Fish., 3, 293; Regan, 1922, Proc. zool. Soc. London, 169. 
Haplochromis pharyngomylus Regan, 1929, Ann. Mag. nat. Hist. (10) 3, 388. 

Description, based on thirty-eight specimens (including the holotype) 70-126 mm. 
S.L. 

Depth of body 33-8-42-0 (M = 38-5) per cent of standard length, length of head 
31-5-36-8 (M = 34-6) per cent. Dorsal head profile straight or somewhat curved 
(occasionally concave between the eyes), sloping fairly steeply. Preorbital depth 
13-8-19-0 (M = 16-8) per cent of head, least interorbital width 23-7-28-5 (M = 26-3), 
length of snout 27-3-33-3 (M = 30-8), diameter of eye 23-0-31-8 (M = 26-5), depth 
of cheek 19-7-27-0 (M = 24-1) per cent of head. Caudal peduncle 13-6-18-5 
(M = 16-2) per cent of standard length, 1-1-1-6 (modal range 1-3-1-5) times as 
long as broad. 

Mouth horizontal or slightly oblique, jaws equal anteriorly, the lower 35-8-44-0 
(M = 38-6) per cent of head, 1-3-2-0 (mode 1-4) times as long as broad. Posterior 
tip of the maxilla reaching or almost reaching the vertical to the anterior orbital 
margin. 

Gill rakers moderately stout, 6-8 (mode 7), rarely 9, on the lower part of the first 
arch, the lowermost 1 or 2 rakers reduced. 

Scales ctenoid ; lateral line with 30 (f.i), 31 (f.6), 32 (f.12), 33 (f.13), 34 (f.4) or 
35 (f.i) scales ; cheek with 3 or 4 series. Six or 7 (less frequently 8) scales between 
the lateral line and the dorsal fin, 7 or 8 (less frequently 6 or 9, rarely 10) between 
the pectoral and pelvic fin bases. 

Fins. Dorsal with 24 (f.7), 25 (f.28) or 26 (f.3) rays, anal with n (f.5), 12 (f.28) 
or 13 (f.5), comprising XV-XVI, 8-10 (rarely n) and III, 8-10 spinous and branched 
rays for the fins respectively. Pectoral 68-5-91-0 (M = 79-6) per cent of head. 
Caudal truncate. 

Teeth. In fishes 70-90 mm. S.L. only unequally bicuspid, relatively stout teeth 
occur in the outer row of both jaws. Fishes 90-100 mm. S.L. show some variability 
in the form of these teeth, which may be unicuspids, a mixture of bi- and weakly 
bicuspid teeth or both bi- and unicuspids. There are 30-42 (mode 36, modal range 
36-40) outer teeth in the upper jaw. 

Teeth in the inner rows are usually tricuspid in fishes less than 100 mm. S.L. In 
larger individuals these teeth may be tricuspid or there can be a mixture of tri- 
and weakly tricuspids ; it is uncommon to find only unicuspid teeth in the inner 
rows. There are 2 or 3 (rarely 4) rows in the upper jaw and 1 or 2 (rarely 3) in the 
lower. 

Lower pharyngeal bone and teeth. With one exception, the lower pharyngeal bone 
is massive, even in the smallest specimens. Nevertheless, a slight size correlated 
increase in relative stoutness can be detected in large fishes. Apart from H. ishmaeli 
(p. 277) and Astatoreochromis alluaudi, no other Lake Victoria Cichlidae have such 
large pharyngeal bones. 



A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 271 

The exceptional specimen mentioned above was caught near Jinja and measured 
98 mm. standard length. The lower pharyngeal of this fish can be compared with 
that of an H. obtusidens of a similar size. Since in all other respects this fish resembles 
H. pharyngomylus more closely than H. obtusidens it is included in the description 
of H. pharyngomylus. Four other specimens with even finer pharyngeal bones and 
dentition, but otherwise resembling H. pharyngomylus, are not included in the 
description but are dealt with in a separate appendix (p. 274). 

The lower pharyngeal teeth of H. pharyngomylus are large and molariform except 
for a few teeth situated in the upper, lateral angles of the bone. These teeth, 
although relatively stout, are cuspidate and small ; the number of such teeth 
decreases markedly in the largest individuals. By analogy with other species having 




Fig. 19. Haplochromis pharyngomylus ; holotype. Drawn by Miss M. Fasken. 



enlarged pharyngeals and for which a greater size range of specimens is available 
is seems likely that small H. pharyngomylus should have less massive bones and 
fewer molariform teeth, confined to the median tooth series (see Greenwood, 1959). 
Osteology of the neurocranium. Apart from an enlarged and strengthened articular 
apophysis for the upper pharyngeals, the neurocranium of H. pharyngomylus is 
that of a large, generalized Haplochromis. The apophysis has been figured and 
described previously (Greenwood, 1954). Compared with the apophysis of H. 
obtusidens that of H. pharyngomylus has a greater surface area and the prootic 
buttresses are more obvious. These characters are clearly correlated with the more 
massive pharyngeal bones and musculature of H. pharyngomylus. Only slight dif- 
rences exist in the apophyseal region of H. pharyngomylus and H. ishmaeli ; but 
both species differ from Astatoreochromis alluaudi in the form taken by the various 
elements contributing to the apophysis (Greenwood, 1959. In that paper, I also 



272 A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 

briefly described the possible effects and interactions of genetical and environmental 
factors on the development of apophyseal form and size in mollusc-crushing species). 

Coloration in live fishes. Adult males. Ground colour blue-grey overlying silver ; 
a distinct coppery sheen on the flanks. Dorsal fin hyaline with pinkish lappets and 
margin to the soft part. Anal hyaline, ocelli yellow. Caudal hyaline with a pink 
flush most intense distally and on the ventral half of the fin. Pelvics black. Females. 
Golden-green, becoming silvery-white ventrally. Dorsal and anal fins dark ; the 
caudal yellowish-green, darker proximally. 

Preserved material. Adult males. Greyish, darker in sexually active individuals ; 
a dark lachrymal stripe sometimes visible ; barring on the flanks variable, but usually 
consisting of seven to ten narrow vertical bars and a very faint longitudinal stripe 
following the course of the upper lateral line. Dorsal fin hyaline, dark basally in 
juvenile and quiescent fishes but almost black in sexually active individuals ; 
lappets dark, the soft dorsal intensely maculate in sexually active fishes. Anal 
hyaline, dark basally, with five or six ocelli arranged in one or two horizontal rows. 




0-2 cm 




0-2 cm 



(a) (b) 

Fig. 20. Haplochromis pharyngomylus ; lower pharyngeal bone, in occlusal view, of 
(a) a fish 98 mm. S.L., and (b) a fish 126 mm. S.L. 

Caudal hyaline in quiescent fishes, dusky in active ones. Pelvics hyaline in juveniles, 
black in adults. Females, silvery-grey to light brown ; striping variable ; when 
present, as described for males. Dorsal fin hyaline or greyish, the soft part weakly 
maculate in adults. Caudal hyaline or dark, the upper half faintly maculate in some 
specimens. Anal and pelvic fins hyaline. 

Distribution. Known only from Lake Victoria. 



Ecology. Habitat. Haplochromis pharyngomylus is apparently confined to water 
less than 40 feet deep and to areas of the lake where the bottom is hard (sand, 
shingle or, less frequently, rock) ; the species is often found amongst stands of 
aquatic plants. A few specimens have been caught in the areas of intergradation 
between sand and mud substrates. 

Food. Analyses of stomach and intestinal contents from thirty specimens (repre- 
senting most localities) show that fishes in the size-range 75-115 mm. S.L. feed 
mainly or even exclusively on Mollusca ; only one fish had eaten mollusca and 
insect larvae. The sample examined also indicated that both bivalves and gastropods 



A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 273 

are eaten in approximately equal proportions. The fragmented shells do not permit 
accurate identification of the prey species ; Sphaerium sp. and Corbicula sp. were 
recognized amongst the bivalve remains and Melanoides tuberculata (the pre- 
dominant snail), Bellamy a and less frequently Biomphalaria amongst the gastropod 
fragments. 

It is clear from the small size of most shell fragments that the pharyngeal mill 
in H. pharyngomylus is an efficient crushing mechanism. Yet, despite this powerful 
barrier some shells do pass into the stomach almost undamaged. Since these shells 
are invariably empty it seems that the digestive enzymes (especially those of the 
stomach) are capable of breaking down the bodies of snails without preliminary 
and physical assistance from the pharyngeal teeth. Trewavas (1938) noticed that 
the gut contents of H. mahagiensis (a mollusc eater from Lake Albert) were " im- 
pregnated and held in a hard mass by botryoidal aggregates of calcite. Whether 
these were formed before or after death and fixation is a matter for conjecture." 
My observations on the gut contents of H. pharyngomylus support Trewavas' 
observations only when the material had been fixed in formol. This would suggest 
that the calcite aggregates are formed as a result of a chemical reaction between 
the slightly acid formol and the calcium of the shell. I have never observed aggre- 
gates in fresh gut contents, where the various shell fragments could be separated 
easily. 

Breeding. The actual spawning sites and breeding behaviour of H. pharyngomylus 
axe unknown, but females carrying embryos and larvae in the buccal cavity occur 
together with non-breeding fishes in most localities. The smallest adult recorded 
is a female 90 mm. S.L. ; the smallest adult male is 94 mm. S.L. Both sexes reach 
the same maximum adult size. 

Affinities and diagnosis. Extreme hypertrophy of the pharyngeal mill serves to 
set H. pharyngomylus apart from all except one species of Lake Victoria Haplo- 
chromis. The other species is H. ishmaeli which appears to be very closely allied to 
H. pharyngomylus. Although in structural characters the two species are similar 
an experienced observer can, in most instances, readily distinguish between the 
two species. But, as is so often the case with Haplochromis, the subjective characters 
used for " field " identifications cannot be quantified or adequately described on 
paper. When seen alive, adult males of the two species have distinctive coloration. 

Haplochromis pharyngomylus is distinguished from H. ishmaeli by the following 
characters : fewer outer teeth in the upper jaw (30-42, modal range 36-40 cf. 38-66, 
modal range 44-52 in H. ishmaeli); shorter pectoral fin (68-0-91-0, M = 79-6 per 
cent of head cf. 75-0-102-0, M = 88-5) ; differences in male breeding coloration (see 
p. 272 and p. 277) and differences in habitat preference (H. pharyngomylus is es- 
sentially a species of hard substrates whilst H. ishmaeli shows a marked preference 
for muddy areas) . With the exception of differences in male breeding colours, none 
of these characters alone is trenchant ; taken together, however, they provide fairly 
reliable diagnostic features. 

The phylogenetic position of H. pharyngomylus has been discussed above (p. 269) 
and in an earlier paper (Greenwood, 1954). 

In Lake Edward, H. placodus Poll represents the equivalent evolutionary phase in 



274 A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 

the development of hypertrophied pharyngeals. Haplochromis placodus and H. 
pharyngomylus could be derived from a common ancestral stem, but of course, the 
two species could equally well be examples of convergent evolution. 

APPENDIX 

Four specimens (115-120 mm. S.L.) from Jinja (B.M. (N.H.) Reg. No. 1959. 
4.28.352-355) bear a strong resemblance to H. pharyngomylus except that the lower 
pharyngeal bone in these specimens is barely enlarged and the pharyngeal dentition 
is correspondingly weak. These fishes cannot be distinguished from H. pharyn- 
gomylus on proportional measurements, teeth of jaws, fin and scale counts, general 
appearance or preserved coloration. All four fishes are adult males but unfortunately 
their live coloration was not recorded. 

From H. pharyngomylus material described above and from field observations on 
numerous other specimens it is clear that intraspecific variability in the enlargement 
of the pharyngeal bones is slight and mainly correlated with size. No specimens 
were found which could be considered intermediate between the typical condition 
for the species and that seen in the aberrant individuals. On the other hand, a 
study of Astatoreochromis alluaudi (a Haplochromis-\ike monotypic genus) showed 
that some populations have enlarged pharyngeals whilst others exhibit only slight 
hypertrophy of these bones (Greenwood, 1959). 

Thus for the moment it is impossible to dismiss the possibility that the four 
" pharyngomylus " -like fishes are indeed aberrant members of that species. I do 
not propose, however, to include them in this revised description or to describe 
them as distinct species until further collections dictate one step or the other. 

Study material and distribution records 



B.M. (N.H. 



nd Reg. No. 


Locality 
Uganda 


Collector 


1906.5.30.383 


Entebbe 


Degen 


1907.5.7-71 


Buddu Coast 


Simon 


1958. 12.5.30-33 


Entebbe 


Pitman 


1959. 4. 28. 317-321 


Beach near Nasu Point 


E.A.F.R.O 


1959.4.28.322-326 


Entebbe, Harbour 


, ,, 


1959.4.28.327-329 


Entebbe, Airport beach 


>> 


1959.4.28.333-334 


Beach near Grant Bay 


,, 


1959.4.28.335-336 


Between Yempita and 
Busiri Isls., Buvuma Channel 


E.A.F.R.O 


1959.4.28.340-343 


Beach near Hannington Bay 


1$ 


1959.4.28.344 


Entebbe, Harbour 


»> 


1959.4-28.345-347 


Napoleon Gulf, near Jinja 


,, 


1959.4.28.351 


Ramafuta Island 


>t 


1959.4.28.358 


Jinja Pier 
Kenya 


" 


1959.4.28.348 


Kasingiri Gingo 
(Kavirondo Gulf) 


»» 



* 



A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 275 

Study material and distribution records (cont.) 

Museum and Reg. No. Locality Collector 

Tanganyika 

1959.4.28.337-339 . Majita 

1959.4.28.349 . Beach near Majita 

1959.4.28.350 . Mwanza, Capri Bay 

Lake Victoria, Locality Unknown 

1928. 5. 24. 313 . — M. Graham 

(Holotype) 

1959.4.28.330-332 . E.A.F.R.O. 

Haplochromis ishmaeli Blgr. 1906 
(Text-fig. 21) 

Haplochromis ishmaeli (part) Boulenger, 1906, Ann. Mag. nat. Hist. (7), 17, 446 ; Idem, 1915, 

Cat. Afr. Fish. 3, 293 ; Regan, 1922, Proc. zool. Soc. London, 169. 
Tilapia pallida (part) Boulenger, 191 1, Ann. Mus. Genova (3), 5, 74; Idem, 1915, torn, cit., 231-2. 
Labrochromis pallidas Regan, 1920, Ann. Mag. nat. Hist. (9), 5, 45 (footnote). 
Tilapia martini (Part) Boulenger, 1915, torn, cit., 239. 
Paratilapia victoriana (part) Boulenger, 1915, torn, cit., 341. 
Haplochromis macrops (part) Regan, 1922, Proc. zool. Soc. London, 166. 

In the original description of H. ishmaeli, Boulenger listed thirteen type specimens 
(syntypes) all collected by Degen at Bunjako, Uganda. However, only eleven 
specimens (ten in spirit and one skeleton) answering this description can be found 
in the collections of the B.M. (N.H.) ; furthermore, only eleven such specimens are 
recorded in the Museum's catalogue of accessions. That the number thirteen was 
a slip of the pen seems certain because Boulenger (1915) only lists ten types from 
Bunjako in the third volume of his Catalogue of Fresh- water Fishes of Africa. The 
only other specimen from this locality is listed in the " Catalogue " as a skeleton 
and is presumably the eleventh syntype. 

These figures agree with the number of specimens I could locate and, more im- 
portant, they agree with the number registered in the Museum's record of accessions. 
But, the situation is still somewhat obscure because in the " Catalogue " (torn. cit. 
loc. cit.) Boulenger lists as a type a specimen collected from Entebbe ; since no 
reference is made to this fish in the original description I am treating its later 
inclusion in the list of types as erroneous. 

Note on the genus Labrochromis, Regan, 1920. This genus, briefly described in a 
footnote to Regan's paper on the genera of Tanganyika cichlids was apparently 
based on a single specimen prepared from one of the syntypes of T. pallida (B.M. 
(N.H.) Reg. No. 1911.3.3.132). In his revision of the Lake Victoria Cichlidae, 
Regan (1922) correctly identified this skeleton as being that of Haplochromis ishmaeli 
and abandoned the genus Labrochromis on the grounds that, apart from possessing 
hypertrophied pharyngeals, H. ishmaeli is " nearly identical with H. cinerens." 
Whilst I do not agree with the latter part of this statement, I fully endorse Regan's 
action in not maintaining the monotypic genus Labrochromis for Haplochromis 
ishmaeli. 

zool. 6, 4 19 



276 A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 

Lectotype of Haplochromis ishmaeli. An adult female 104-0 + 23-0 mm. total 
length (B.M. (N.H.) Reg. No. 1906.5.30.400) collected by Degen at Bunjako, 
Uganda. 

Description, based on thirty-five specimens (including the lectotype and two 
paratypes) 82-136 mm. S.L., all from Lake Victoria. Two specimens from Lake 
Edward are described separately on p. 278. 

Depth of body 37-0-45-5 (M = 40-1) per cent of standard length, length of head 
33-8-37-5 (M = 34*8) per cent. Dorsal head profile slightly curved or straight, 
sloping fairly steeply. Preorbital depth 15-3-20-5 (M = 17-0) per cent of head, 
least interorbital width 24-0-32-0 (M = 27-6), length of snout 29-0-36-0 (M = 31-6), 
diameter of eye 23-0-31-0 (M = 27-7), depth of cheek 20-7-31-0 (M = 25-5) per 
cent. Caudal peduncle 14-6-18-8 (M = 17-6) per cent of standard length, 1-2-1-6 




Fig. 21. Haplochromis ishmaeli ; lectotype (from Boulcnger, Fishes of the Nile). 



(modal range 1-3-1-5) times as long as deep. Jaws equal anteriorly, the lower 
35-8-42-5 (M = 39-1) per cent of head and 1-4-2-0 (modal range 1-4-1-6) times as 
long as broad. Mouth horizontal or very slightly oblique ; the posterior tip of the 
maxilla reaching the vertical through the anterior orbital margin or somewhat 
beyond, rarely not quite reaching the orbit. 

Gill rakers stout, but relatively slender in a few specimens ; 6-8 (mode 7), rarely 
9, on the lower part of the first arch, the lowermost one or two rakers usually reduced. 

Scales ctenoid; lateral line with 30 (f.3), 31 (f.8), 32 (f.15), 33 (f.5) or 34 (f.4) 
scales ; cheek with 3 or 4 rows. Five to 7 (rarely 8) scales between the lateral line 
and the dorsal fin origin ; 8-9 (occasionally 7) between the pectoral and pelvic 
fin bases. 

Fins. Dorsal with 23 (f.i), 24 (f.13) or 25 (f.21) rays, anal with 11 (f.13), 12 (f.21) 
or 13 (f.i) comprising XV-XVI, 8-10 and III, 8-10 spinous and branched rays 
for the fins respectively. Pectoral fin 75-0-102-0 (M = 88-5) per cent of head. 
Caudal truncate or subtruncate. 

Teeth. The outer row in both jaws usually contains a mixture of relatively stout, 
unequally bicuspid and unicuspid teeth ; less frequently only unicuspids occur in 



A REVISION OF THE LAKE VICTORIA HAPLOCHROMJS SPECIES 277 

this row. There is apparently no size-correlated difference in the type or the number 
of teeth present. There are 38-66 (modal range 44-52) teeth in the outer row of 
the upper jaw. 

Teeth in the inner series are generally tricuspid, but in some fishes the entire 
inner series are composed of unicuspids. There are 2 or 3 (rarely 1) rows of teeth 
in the upper jaw and 1 or 2 (less commonly 3) rows in the lower. The inner teeth 
are implanted somewhat obliquely. 

Lower pharyngeal bone and teeth. The lower pharyngeal bone in H. ishmaeli is 
massive and almost all the teeth are molariform ; only those situated in the posterior 
angles of the bone remain small, with pointed crowns. There is some individual 
variation in the relative enlargement of the bone, but this is slight in comparison 
with the variation known from such species as H. obtusidens and Astatoreochromis 
alluaudi. A slight size-correlated increase in the relative stoutness of the pharyngeal 
bones was observed in the material studied ; likewise, there is an increase in the 
number of molariform teeth in larger fishes. 

In size, shape and dentition, the pharyngeal bones of H. ishmaeli are directly 
comparable with those of H. pharyngomylus. 

Neurocranial osteology. The neurocranium of H. ishmaeli is virtually identical 
with that of H. pharyngomylus. 

Coloration of live fishes. Sexually active males. Ground colour light yellow-green 
dorsally, shading to yellow on the flanks and greyish-white ventrally. Dorsal fin 
yellow-green, lappets of the anterior spines dusky, the remainder scarlet, as are the 
spots and dashes between the branched rays. Caudal fin greyish, with red maculae 
(sometimes coalesced) between the rays. Anal smoky grey with black lappets and 
an overall scarlet flush ; ocelli yellow. Pelvic fins black. Females golden-green 
shading to silvery-white ventrally, the pectoral region faintly blackish. Dorsal 
fin hyaline, with a narrow red margin. Anal light yellowish-green. Caudal and 
pelvic fins pale yellow. 

Preserved material. Adult males, yellowish-grey, dusky on the chest and branchio- 
stegal membrane (lighter in sexually quiescent fishes) ; very faint indications of 
six or seven vertical stripes on the flanks. (These bars are more widely spaced than 
in H. pharyngomylus, see p. 272). A distinct lachrymal stripe. Dorsal fin hyaline 
in juvenile fishes but darker and with the soft part maculate in mature males ; 
lappets dark. Caudal hyaline, the upper part often maculate. Anal hyaline, the 
basal third dark in sexually active individuals ; three to five large, dead-white 
ocelli arranged in one or, rarely, two rows. Females, brownish to yellowish-silver ; 
sometimes, very faint traces of five to seven transverse bars on the flanks ; a weak 
lachrymal stripe visible in some fishes. All fins hyaline, the dorsal darkest. 

Distribution. Lakes Victoria and Edward. 

Ecology. The data given in this section relate to fishes from Lake Victoria ; 
nothing is known for fishes from Lake Edward. 

Habitat. Haplochromis ishmaeli is essentially a species of inshore regions where 
the water is less than 30 feet deep and the bottom composed of soft, organic mud. 



278 A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 

No specimens have been recorded from depths greater than 60 feet, but some have 
been caught in nets set over sand and shingle substrates. Haplochromis ishmaeli 
would seem to be the ecological (soft substrate) counterpart of H. pharyngomylus 
(solid substrates). 

Because of the difficulty in distinguishing H. ishmaeli from H. pharyngomylus and 
especially because the latter species was not recognized until after Graham's col- 
lections were brought to England, it is impossible to use Graham's (1929) catch 
records as an additional source of information on the intralacustrine distribution of 
H. ishmaeli. 

Food. The stomach and intestinal contents of nineteen fishes (from several 
localities) indicate that H. ishmaeli in the size-range 84-135 mm. S.L. feed almost 
exclusively on Mollusca ; the few insect larvae found together with snails in the 
stomach of one fish suggest that insects may be ingested accidentally. 

The number of fishes with food remains is insufficient to determine whether 
bivalves or gastropods predominate in the diet. Slightly more gastropods (Melanoides 
tuberculata) than bivalves (Corbicula sp.) were found in the sample examined. 

Breeding. No information is available on the breeding sites or habits of H. ishmaeli 
although the number of females with " spent " ovaries and ventrally distended 
mouths suggests that the species is probably a female mouth-brooder. The smallest 
mature females are 97 mm. S.L. (but see Appendix 2) and the smallest adult male 
is 98 mm. S.L. Females apparently reach a greater maximum size than do males. 

Affinities and diagnosis of H. ishmaeli are discussed on page 273, with reference 
to its closest relative, H. pharyngomylus. 

APPENDIX 

(1) Haplochromis ishmaeli from Lake Edward. I have been able to examine only 
two specimens from Lake Edward, one an adult female 120-0 + 28-0 mm. long and 
the other an adult male n8-o -j- 27-0 mm. long (see Trewavas, 1933). In general 
appearance, in most morphometric characters and in scale and fin ray counts the 
two fishes are indistinguishable from specimens of a similar size from Lake Victoria. 
The nature of the pharyngeal bones and dentition is also identical. Nevertheless, 
the Lake Edward fishes do differ slightly in three characters. 

(i) The interorbital is somewhat narrower (24-1 and 25-0 per cent of head) than 
the mean interorbital width of Victoria fishes (27-6 per cent) although still within 
the range known from this population, (ii) The number of outer teeth in the upper 
jaw (40 and 42) is in the lower section of the range for Lake Victoria fishes, (iii) The 
caudal peduncle is stouter in the majority of Victoria specimens. Now that more 
specimens are available from Lake Victoria, Trewavas' (op. cit.) observation on the 
larger eye of the Edward specimens is no longer applicable. 

None of the differences commented upon above is so marked as those charac- 
terizing the Lake Victoria and Lake Edward populations of Astatoreochromis alluaudi 
(Greenwood, 1959). 

Nothing is known about the bionomics of H. ishmaeli in Lake Edward ; the two 
specimens studied were caught in a seine net fished from the eastern shore of the 
lake near Kisenyi. 



A REVISION OF THE LAKE VICTORIA H A PLOCHROMIS SPECIES 



279 



(2) Three H. ishmaeli-like fishes from Lake Victoria. Three specimens (82, 96 and 
100 mm. S.L. (B.M. (N.H.) Reg. Nos. 1959.4.28.304-306)) caught in a trawl off 
the mouth of the Nzoia river, Kenya, present something of a problem. In appearance 
and in all standard counts and measurements these fishes are typically H. ishmaeli. 
However, their collector, Mr. S. H. Deathe, recorded the live coloration of these 
fishes as bright pink. Because all three specimens are adult males, I consider that 
this striking departure from the usual male coloration of H. ishmaeli may be 
significant, especially since pink is not one of the basic colours in H. ishmaeli 
pigmentation. A further interesting difference is that the smallest specimen is 
sexually mature whereas the smallest adult H. ishmaeli recorded is 98 mm. S.L. 

Unfortunately I have only these three specimens and I did not see them when 
alive. Thus, it is difficult for me to assess fully this seemingly outstanding difference 
in coloration. 

No typical H. ishmaeli were reported in the same haul and I do not consider 
that there is enough evidence to decide whether these peculiar individuals represent 
an aberrant population of H. ishmaeli or a distinct species differing from H. ishmaeli 
in the coloration of its adult males. 



Study material and distribution records 



Museum and Reg. No. 



Locality 



Collector 



Uganda 



.M. (N.H.). — 1906.5.30.400 


Bunjako 


Degen 


(Lectotype) 






1906.5.30.401-402^ . 


,, 


,, 


(Paratypes) 






1906.5.30.396 


,, 


,, 


(Paratype) 






1911.3.3.131 


Jinja 


Bayon 


(Paratype Tilapia pallida) . 






1909. 5. 4. 8-10 


Sesse Isls. 


>> 


1906.5.30.275-279 . 


Bunjako 


Degen 


1959. 4. 28. 281-290 


Pilkington Bay 


E.A.F.R.O. 


1959.4.28.296-300 


Ekunu Bay 




1959.4.28. 301-303 . 


Pilkington Bay 




1959.4.28.307-309 . 


Buka Bay 




i959-4- 28 -3i2-3i3 . 


Entebbe, Harbour 




1959. 4. 28. 314 


Macdonald Bay 




1959. 4. 28. 316 


o°4'S„ 33 14' E. 
Kenya 




1909. 11 . 15.40 


Kisumu 


A. B. Percival 


1959.4.28.274-280 


South of Port Southby 


E.A.F.R.O. 


1959-4-28.356 


Kavirondo Gulf 

Tanganyika 


M. Graham 


1959. 4. 28. 291-295 . 


Majita 


E.A.F.R.O. 


1959.4.28.310-311 . 


Beach near Majita 


• 



2 8o A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 

SUMMARY 

1. Eleven species are redescribed on the basis of new material. 

2. In addition, three new species [Haplochromis theliodon ; H. empodisma and 
H. saxicola) are described. 

3. Several phyletic lines are represented amongst these fourteen species, which 
include H. cinereus, a species once thought to represent one of the basic types from 
which the present-day species-flock had evolved. Evidence now available suggests 
that, anatomically, H. cinereus is not sufficiently generalized to retain this distinction. 

4. Notes are given on the feeding habits and bionomics of the species. 

5. Three species (H. obtusidens, H. pharyngomylus and H. ishmaeli) are largely or 
entirely mollusc-eaters ; three others (H. humilior, H. theliodon and H. riponianas) 
feed on both insects and molluscs ; one, H. martini, is a piscivorous predator and 
seven others are insectivore/omnivores. 

6. One species, H. niloticus (nom. nov. for Tilapia bayoni Blgr. 191 1) is known 
only from the Victoria Nile, whilst H. ishmaeli occurs in both Lakes Victoria and 
Edward ; H. humilior is found in Lake Victoria and the Victoria Nile. 

7. The assumed distribution of H. macrops in both Lakes Victoria and Edward is 
discussed ; no definite conclusion can be drawn from the material now available. 

8. Two groups of specimens are dealt with in separate appendices because of their 
uncertain taxonomic position. One group is apparently allied to or even conspecific 
with H. pharyngomylus and the other with H. ishmaeli. No conclusion can be 
reached on the status of these aberrant fishes. 



ACKNOWLEDGMENTS 

It is with great pleasure that I acknowledge my thanks to Dr. Ethelwynn Trewavas 
for her advice and criticism and for reading the manuscript of this paper. My thanks 
are also due to Mr. C. C. Cridland of E.A.F.R.O. Jinja, Uganda, for identifying 
many of the Mollusca recovered from gut contents. 



REFERENCES 

Brooks, J. L. 1950. Speciation in ancient lakes. Quart. Rev. Biol. 25 : 131. 

Graham, M. 1929. A report on the Fishing Survey of Lake Victoria, 1927-1928, and Appendices. 

Crown Agents, London. 
Greenwood, P. H. 195 1. Evolution of the African Cichlid fishes ; the Haplochromis species 

flock in Lake Victoria. Nature, London, 167 : 19. 

1954- O n two cichlid fishes from the Malagarazi River (Tanganyika), etc., Ann. Mag. nat. 

Hist. (12), 7 : 401. 

1956a. The monotypic genera of cichlid fishes in Lake Victoria. Bull. Br. Mus. nat. 

Hist., Zool. 3 : No. 7. 

1 956ft. A revision of the Lake Victoria Haplochromis species (Pisces, Cichlidae). Part I. 

Ibid, 4 : No. 5. 

1957- A revision of the Lake Victoria Haplochromis species, etc. Part II. Ibid. 5 : No. 4. 

1959- The monotypic genera of cichlid fishes in Lake Victoria, Part II, and A revision 

of the Lake Victoria Haplochromis species, etc. Part III. Ibid. 5 : No. 7. 



A REVISION OF THE LAKE VICTORIA HAPLOCHROM I S SPECIES 281 

Pappenheim, P. & Boulenger, G. A. 1914. Fische. Wissenschaftliche Ergebnisse der 

Deutschen Zentral-Afrika Expedition, 1907-1908, 5 : 225. 
Poll, M. 1939. Poissons. Exploration du Pare National Albert, mission G. F. de Witte 

(i933- I 935)> fasc - 2 : 7 8 - 
& Dam as, H. 1939. Poissons. Exploration du Pare National Albert, mission H. Damas 

(i935~ I 93 6 ). fasc - 6 : 73- 
Regan, C. T. 1921. The cichlid fishes of Lakes Albert Edward and Kivu. Ann. Mag. nat. 

Hist. (9), 8 : 632. 
Trewavas, E. 1933. Scientific results of the Cambridge expedition to the East African 

lakes, 1930-1931. II. The cichlid fishes. /. Linn. Soc. (Zool.), 38 : 309. 
1938. Lake Albert fishes of the genus Haplochromis. Ann. Mag. nat. Hist. (11), 1 : 435. 




ON A STYGIOMYSIS FROM THE 

WEST INDIES, WITH A 

NOTE ON SPELAEOGRIPHUS 

(CRUSTACEA, PERACARIDA) 



ISABELLA GORDON 




BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ZOOLOGY Vol. 6 No. 5 

LONDON: i960 



ON A STYGIOMYSIS FROM THE WEST INDIES, 

WITH A NOTE ON SPELAEOGRIPHUS 

(CRUSTACEA, PERACARIDA) 



BY 



ISABELLA GORDON, D.Sc, Ph.D. 



Pp. 283-324. 
With 1 Table, Plates 3-4 and 34 Text-figures. 




BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ZOOLOGY Vol. 6 No. 5 

LONDON : i960 



THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted in 1949, is 
issued in five series, corresponding to the Departments 
of the Museum, and an Historical series. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

This paper is Vol. 6, No. 5 of the Zoological series. 



Trustees of the British Museum, i960 



PRINTED BY ORDER OF THE TRUSTEES OF 
THE BRITISH MUSEUM 

Issued March, i960 Price Fifteen Shillings 



ON A STYGIOMYSIS FROM THE WEST INDIES : 
WITH A NOTE ON SPELAEOGRIPHUS 
CRUSTACEA, PERACARIDA; 

By ISABELLA GORDON, D.Sc, Ph.D. 



SYNOPSIS 
Stygiomysis holthuisi (Gordon), the second known species of the aberrant Mysidacean genus 
Stygiomysis, is described and figured in detail. The systematic position of the family Stygio- 
mysidae is discussed. It is most closely allied to the family Lepidopidae and both families possess 
certain characters of the suborder Mysida, others of the suborder Lophogastrida. The para- 
gnatha of some other Mysidacea are figured for comparison with those of Stygiomysis. The termin- 
ology of the thoracic limbs, and of the segments of the thoracic endopodite, is briefly discussed. 
A note on Spelaeogriphus lepidops Gordon is added, together with a few comments on the 
systematic position of the order Spelaeogriphacea. 



CONTENTS 

I. On Stygiomysis holthuisi (Gordon) from the West Indies : 
Introduction ....... 

Diagnosis of the genus Stygiomysis 
Description of Stygiomysis holthuisi (Gordon) 
II. On the Systematic Position of the Stygiomysidae 
Diagnosis of the family Lepidopidae 

III. Note on the Paragnatha in Some Mysidacea 

IV. The Terminology of : — 

(a) the thoracic appendages ..... 

(b) the segments of the thoracic endopodite 
V. Additional Note on Spelaeogriphus lepidops Gordon 



Page 

285 
286 
287 
299 
308 
309 

315 
319 
320 



I. ON STYGIOMYSIS HOLTHU ISI (GORDON) 
FROM THE WEST INDIES. 

INTRODUCTION. 

Dr. L. B. Holthuis of the Rijksmuseum van Natuurlijkc Historie, Leiden, kindly 
sent me for study three specimens of an interesting cavernicolous " shrimp " which 
he collected in February, 1957 on the island of St. Martin, Lesser Antilles. When I 
first examined these specimens I thought that, in spite of the long vermiform body 
and very short carapace, they looked remarkably like Mysidacea. Then, when I 
discovered some of their most striking peculiarities, I decided that they did not 
belong to the Order Mysidacea as at present defined. I fully intended to look up 
all the literature relating to cavernicolous Mysidacea and, had I done so, I certainly 
would not have referred these specimens to a new genus. But, when I wrote the 
preliminary note to Nature (Gordon, 1958), I did not realise that I had inadvertently 



ZOOL 6, 5 



20§ 



286 ON A STYGIOMYSIS FROM THE WEST INDIES 

omitted to refer to Caroli's 1937 paper on the genus Stygiomysis. I have to thank 
Dr. H. Nouvel of Toulouse for calling my attention to this lapse and I fully agree 
with him that my genus Rhopalonums is a synonym of Stygiomysis Caroli. 

The discovery of a second species of Stygiomysis so far from the locality of the 
type species of the genus, which is in Italy, is very interesting. Perhaps this rare 
genus is, or was at some remote period, widely distributed. Caroli had only four 
specimens of Stygiomysis hydruntina when he published his " nota preliminaire " 
in 1937. As far as I know, he has never published a fuller description of the species, 
nor has it been collected again although the two Italian grottos must in all proba- 
bility have been visited since 1937. I made a fairly complete study of the three 
type specimens of Stygiomysis holthuisi and had finished the description and figures 
before I read Caroli's Stygiomysis paper. By that time I also had come to the con- 
clusion that the species was in fact an aberrant member of the Mysidacea, thus 
confirming Caroli's opinion. So far no ovigerous female of either species has been 
found, although in each case a somewhat immature female with only four pairs of 
developing oostegites is known. Caroli thought it most probable that in Stygiomysis 
the full complement of oostegites will be seven pairs ; within the order Mysidacea 
there are as a rule either seven pairs of oostegites, or else two or three pairs — Thalas- 
somysis established in 1939 has four pairs, the first two of which are small, the last 
two pairs forming most of the brood pouch. But, as Caroli rightly points out, in 
no Mysid are oostegites absent from the last pair of thoracic limbs. If Stygiomysis 
should prove to have only four pairs, then they would be absent from the last pair 
of thoracic appendages and quite exceptional. 

Recently Dr. Holthuis sent me another specimen from the type locality ; this 
proved to be a male and the second pair of pleopods differs appreciably from that 
of the female. Caroli says tkat in Stygiomysis hydruntina none of the pleopods 
" mostra particolari modificazione sessuali ". This admittedly rather slight sexual 
dimorphism of the second pair of pleopods is similar to that found in all three 
monotypic genera of the cavernicolous family Lepidopidae (previously Lepidoph- 
thalmidae), to which the family Stygiomysidae appears to be most nearly related. 

After the manuscript was finished, I received from the Amsterdam Museum a 
fifth specimen which Dr. J. H. Stock obtained in the same place on St. Martin on 
4th February, 1959. 

Order MYSIDACEA 
Genus STYGIOMYSIS Caroli 1937 

Stygiomysis Caroli, 1937, P- 22 °- 
Rhopalonums Gordon, 1958, p. 1552. 

Diagnosis. Aberrant cavernicolous Mysidacea with body rather vermiform, 
somewhat depressed. Carapace short, smooth, fused with tergites of first four 
thoracic somites, not produced antero-laterally beyond the minute rostral apex 
and lacking the usual postero-lateral " wings " so that thoracic somites 5 to 8 are 
entirely exposed. Division of the long thorax into subtagmata further emphasized 



ON A STYGIOMYSTS FROM THE WEST INDIES 287 

by the modification of the first four pairs of thoracic limbs to assist in feeding (maxil- 
liped and gnathopods — see p. 315), the last four pairs being slender peraeopods. 
Abdomen widest across third somite, sixth somite shorter than fifth. Telson almost 
as wide as long ; spines on truncate posterior margin only. Ocular scales broader 
than long, without trace of pigment or visual elements. Antennula more robust 
and rather longer than antenna, the inner flagellum slightly longer than carapace. 
Antennal scale minute ; both segments of antennal protopodite distinct (peduncle 
therefore four-segmented). Labrum broadly triangular. Labium with long, pendant, 
widely separated paragnatha which conceal part of mandibles. Mandibles with well 
developed incisior, poorly developed molar, process and a distinct spine-row ; 
a number of long simple spines on minute third segment of the palp. Maxillula 
without reflexed palp, distal endite unusually large and robust, the anterior apical 
spine very large. Maxilla beset with strong spines rather than setae ; the two 
segments of endopodite very short. Maxilliped (first thoracic limb) pediform, without 
endites or lobes ; exopodite a small unsegmented lamella ; epipodite rather short 
and broad. Thoracic appendages 2 to 8 each with well developed multiarticulate 
exopodite ; endopodites 2 to 4 modified as gnathopods, forwardly directed and 
stronger than those of the peraeopods (5 to 8). Penultimate segment of thoracic 
endopodites not subdivided. Pleopods all reduced, biramous, with one-segmented 
endopodite and three-segmented exopodite (exopodite two-segmented in male 
pleopod 2) ; alike in both sexes, apart from the sexual dimorphism exhibited by 
pleopods 2 in Stygiomysis holthuisi. A transverse chitinous lamella, or pair of 
lamellae, with which the protopodites of the pleopods are incorporated, projects 
from the posterior sternal margin of abdominal somites 3 to 5. Uropods unusual 
in having a long, stout, backward prolongation of the protopodite, armed with 
strong curved spines ; statocyst absent. Oostegites at least four pairs, but the full 
complement is unknown (Caroli thinks it may be seven pairs) . 

Type species : Stygiomysis hydruntina Caroli 1937, from Italy. A second species 
from the West Indies is described below. 



Stygiomysis holthuisi (Gordon) 
Rhopalonuncs holthuisi Gordon, 1958, p. 1552. 

Material. The holotype, a female with developing oostegites measuring 9 mm. 
in length ; the larger paratype is a slightly immature female without oostegites, 
the smaller paratype is probably immature. 1 In addition to the three original 
specimens, I received from the same locality at a later date, a male measuring 
10 mm. in body length (also in the Leiden Museum). 

Locality. Devil's Hole, near Simson's Bay (or Simpson's Bay), in the Dutch 
part of St. Martin, Lesser Antilles, 10th February, 1957, (the three types) and, 
25th May, 1958 (one male). 

1 The holotype and smaller paratype are in the Leiden Museum Collection, and are registered as 
Crust. G. 434 and Crust. G. 435 respectively. The larger paratype is in the British Museum (reg. no. 
1959. vii. 3.1). 



288 ON A STYGIOMYSIS FROM THE WEST INDIES 

Habitat. " Devil's Hole lies to the south east of Simson's Bay bridge near the 
southern shore of St. Martin, just east of Simson's Bay. It is a sink-hole about 5 m. 
deep in a limestone slope. The hole itself is wide and rather easily accessible. From 
the hole a few short dead-ending passages run inwards. In one of these passages, 
which is about 10 m. long, there is a small puddle, about 30 cm. in diameter and 10 
to 20 cm. deep. A film of limestone particles covered the larger part of the surface, 
and a limestone sediment on the bottom made the water turbid when stirred. The 
water tasted saltish, its salinity being 8100 mg. Cl/litre. It was not in total darkness 
as the entrance of the passage could still be seen. The distance of the sink-hole 
from the sea is about 300 m." 

Dr. Holthuis returned to the same pool two days later, but did not obtain any 
specimens of Stygiomysis holthuisi, although he got a few Amphipods which he had 
not seen during his first visit to the cave. Dr. P. Wagenaar Hummelinck, who 
collected in the same locality on 26th July, 1955, obtained only Amphipods and 
Nematodes. (Information supplied by Dr. L. B. Holthuis.) 

Additional material. A fifth specimen was received after the manuscript 
was finished. It was found in a small hole or pitch a few inches square, in the type 
locality, Devil's Hole, St. Martin, by Dr. J. H. Stock on 4th February, 1959. It is 
a slightly immature specimen, probably female, without oostegites, and is in the 
Amsterdam Museum. 

Description of female. 

The largest specimen, measuring 9 mm. in length, and selected as the holotype, 
is represented in Plate 3. The long, narrow, rather vermiform body is flattened 
dorsoventrally and is fully segmented behind the carapace, which is just over one- 
fourth of the total length. The ratio of length to maximum width of carapace is 
i-6 : 1 and the surface is smooth (with perhaps the merest hint of the cervical 
furrow). The carapace is completely fused dorsally with the tergites of thoracic 
somites 1 to 4, being free only at the sides to enclose long narrow branchial chambers. 
The anterior margin, on either side of the median triangular apex, which scarcely 
merits the name of rostrum is rather oblique and slightly concave. In front of each 
concavity is a short broad scale, presumably an ocular scale without any trace of pig- 
ment or visual elements, ((PL 3 and Text -figs. 1-3, 0). This scale curves downwards 
and inwards as shown in Text-figs. 2 and 3. The posterior margin of the carapace 
does not overlap any part of the fifth (first free) thoracic somite ; indeed it is slightly 
overlapped by the latter in the mid-dorsal line (Text-fig. 1 and PI. 3, 5). 

The four free thoracic somites pass imperceptibly into the abdomen, although the 
two tagmata can easily be recognized by their respective appendages. Together 
they form a very elongated barrel (in dorsal aspect) , widest across abdominal somites 
2 and 3 ; abdominal somites 4 and 5 are longest and equal. The telson is as long as 
the fifth and sixth abdominal somites together, and is nearly as broad as long. The 
lateral margins are only slightly convergent and the wide posterior margin is armed 
with strong spines namely, three at each side and a median one flanked by a pair 
of shorter spines ; four or five spinules fill the gaps between lateral and median 
spine groups (PL 3 ; only the median group is visible in Text-fig. 8). 



ON A STYGIOMYSIS FROM THE WEST INDIES 289 

The antennula is longer, and much stouter, than the antenna and is biramous 
(Text-fig. 1 and PL 3). Of the three segments in the protopodite the second is the 
longest, the first or basal one the widest. The flagella are only slightly unequal in 
length, the inner one is the more slender, with about 28 segments whereas the outer 




0.5 mm 



in lateral aspect, after removal of the mouthparts and some thoracic appendages. Fig. 
2. Antero-lateral angle of carapace, basal segment of antennula and antennal peduncle, 
part of labrum. Fig. 3. Fdght anterior portion of body, in ventral aspect. 

/. Paragnath of lower lip. m. Mandible. 0. Ocular scale. 5. Antennal scale, u. Upper 
lip or labrum. 5. Fifth (first free) thoracic somite. 1. First abdominal somite. 

ZOOL 6, 5 



2 9 o ON A STYGIOMYSIS FROM THE WEST INDIES 

one has only 16 to 18 longer segments most of them with long sensory hairs or 
aesthetascs on the inner margin. The antennulae arise beneath the ocular scales 
and are not widely separated. 

The antenna is shorter and much more slender than the antennula. The peduncle 
of the right antenna of the larger paratype is represented, highly magnified, in 
Text -fig. 2. The protopodite seems to have only two segments ; the basal one is 
nearly twice as long as wide, the second one is short and bears, at its outer distal 
angle, a minute exopodite or scale (s). The two basal segments of the endopodite 
are each long and slender (at least four times as long as wide) and the flagellum, 
which has 16 to 18 segments, is about four-fifths of the carapace length (PL 3). 

Mouthparts. Those on the right side of the larger paratype were removed for 
detailed examination. Each appendage was sketched from various aspects before 
any attempt was made to stain and mount it. In this way any distortion due to 
movement of, or pressure from, the coverslip was eliminated. Certain details not 
apparent in the unstained appendages were inserted later from the micro-prepara- 
tions. 

The labrum or upper lip is large and triangular, with rounded apex, beyond which, 
at a lower level, a small rounded lobule is visible (i.e. the lobule is more dorsal in 
position) — see Text-figs. 1 and 3, u. Behind the mandible is the labium or lower 
lip which is unusual in that the paragnatha are widely separated ; moreover, each 
is enormous, pendant and forwardly directed so as to overlap and partially conceal 
the mandible (Text-figs. 1 and 3, /). The right mandible is shown in situ in Text-fig. 3 
and the distal portion is represented in Text-fig. 4. The crown consists of a well 
developed, three-lobed incisor process (with a subsidiary tooth at the base of one 
of the lateral ones) a small but well chitinized knob-like molar process and, between 
the two processes, a strongly chitinized triangular lobef and a row of five graded 
spines. The palp has the usual three segments ; the median one is very long and 
somewhat twisted, the short terminal one has seven long spines. The maxillula 
is unusually large and massive (Text-fig. 5). The proximal endite, Hansen's lobe 1, 
has four terminal spines, two long and plumose, two short and simple. The distal 
or major endite, Hansen's lobe 3, has a terminal row of spines the anterior one 
unusually long and stout and, near its base, is a long seta. There is no backwardly 
directed endopodite or palp. The maxilla is a thin leaf -like appendage abundantly 
beset with spines and setae (Text-fig. 6). Hansen's pi. 3 was not apparent until the 
appendage had been stained and mounted. The proximal endite (s), in this case the 
lobe from 2, is simple, with a median seta on the posterior margin and a series of 
apical and subapical spines. The endite from pi. 3 (d) is divided into a small distal 
lobe with four terminal spines, and a large oval proximal lobe with a row of nine 
marginal spines. A short, two-segmented endopodite (en) follows ; the distal segment 
bears three long spines, the proximal one has two spines on the inner margin. A long 
narrow exopodite (ex) arises from the outer margin of pi. 3 ; its margins, the free 
inner one excepted, bear plumose setae. 

The first thoracic appendage or maxilliped is represented in Text-fig. 7 at a lower 
magnification than the other mouth parts. Though much smaller than, it is essenti- 

f The fixed lacinia, against which the lacinia mobilis works. 



ON A STYGIOMYSIS FROM THE WEST INDIES 



291 



ally similar to, the three following pairs of thoracic limbs. However, it alone bears 
a fairly large epipodite reflexed into the brancial chamber and, moreover, the exopo- 
dite is reduced to a small oval unsegmented lamella (which may also be respiratory). 




Figs. 4-6. Stygiomysis holthuisi (Gordon). Mouthparts of larger paratype. Fig. 4. 
Distal portion of right mandible, with palp. Fig. 5. Right maxillula. Fig. 6. Right 
maxilla. All at higher scale except upper Fig. 4. 

d. Divided endite. en. Endopodite. ex. Exopodite. s. Simple or undivided endite. 
2 and 3 Hansen's plates 2 and 3 of protopodite (coxa and basis). 



292 



ON A STYGIOMYSIS FROM THE WEST INDIES 



The endopodite — a continuation of the axis of the protopodite — comprises five 
segments and a strong terminal claw ; the " knee " joint between the third and fourth 
segments is indicated by an arrow. There is a row of strong spines on the inner 
margin of segments 2 and 3, a group of spines on the distal half of the inner surface 
of segment 4 and a pair of claw-like spines, flanking the terminal claw, on segment 5. 
(Since the terminology of the segments of the endopodite varies, I have thought it 
better in the description of the thoracic limbs to use numbers — see p. 319 for discus- 
sion of this). 




Figs. 7 and 8. Stygiomysis holthuisi (Gordon). Fig. 7. Right first thoracic limb 
(maxilliped) of larger para type. Fig. 8. Posterior part of body of holotype, in ventral 
aspect, e. Exopodite. b. Basis, ep. Epipodite. m. Membranous free lamina connecting 
and incorporating protopodites of pleopods 5. t. Telson. p. Backward prolongation of 
protopodite of uropod. 5 and 6. Fifth and sixth abdominal somites. 3 and 4. Segments 
of endopodite on either side of " knee " joint (indicated by arrow). 



Thoracic appendages 2 to 8. Each of these bears a well developed, natatory exopo- 
dite, as in most Mysidacea. All are visible in dorsal aspect, as represented in PL 3, 
each is composed of a long robust proximal segment followed by a more slender 
tapering flagellum of 10 to 12 segments (Text-figs. 9, 10, n, e). Each segment of 
the flagellum carries a pair of plumose setae, but only the base of each is inserted 
in Text-figs. 9-1 1. 

These seven pairs of appendages are arranged in two distinct groups. The three 
anterior pairs have their endopodites directed forwards, more robust than, and obvi- 



OX A STYGIOMYSIS FROM THE WEST INDIES 



293 




Figs. 9-1 1. Stygiomysis holthuisi (Gordon). Larger paratype. Fig. 9. Right second 
thoracic limb (first gnathopod). Fig. 10. Fourth thoracic limb. Fig. 11. Eighth thoracic 
limb. 

b. Basis, e. Exopodite. 1 to 6. Segments of endopodite. " Knee " joint indicated 
by arrow. 



294 ON A STYGIOMY SIS FROM THE WEST INDIES 

ously differing in function from, those of the four posterior pairs which are slender 
ambulatories. The three anterior pairs should be termed gnathopods since they are 
probably used to seize prey and pass it to the mouthparts. Thoracic appendages 
2, 4 and 8 were removed from the right side of the larger paratype and are repre- 
sented in Text-figs. 9, 10, and 11 respectively. Appendage 2 is, however, represented 
in inner, the other two in outer, aspect. The endopodite, beyond the basis which 
also bears the exopodite (e), comprises, as already mentioned in connection with the 
maxilliped, five segments, or six if the terminal claw is included (Text-fig. n, 1-6). 
The endopodite of gnathopod 1 (thoracic appendage 2) is very robust ; segments 1 
and 2 are short, 3 is the longest and by far the broadest, 4 is long and armed with a 
longitudinal series of six setae on the inner surface and 3 on the ventral margin 
(with another near the margin) , segment 5 is not quite half as long as 4 and is armed 
with a number of short spines in addition to the terminal claw. Gnathopod 2 
(appendage 3) is very similar to, but a trifle less robust than, gnathopod 1 and there 
are only 4 long setae on the distal half of the inner surface of segment 4. As illustrated 
in Text-fig. 10, the third gnathopod is much more slender than gnathopod I, with 
only three setae on the distal third of the inner surface of segment 4. Thoracic 
appendages 4 to 8 are all very similar ; they are in fact peraeopods or walking legs, 
with the endopodite slender as represented in Text-fig. 11. Segment 4 is the longest 
but segment 5 and the terminal claw are also elongated so that the portion distal to 
the " knee " is longer than the proximal part. On the ventral margin of segment 5 
is a row of curved serrated spines. 

The pleopods are similar in form and do not differ much in size from the first to 
the fifth pairs. Each is a small, biramous appendage comprising a short protopodite, 
a one-segmented endopodite and a three-segmented exopodite (Text-figs. 8, 14 and 
16). The first pair are more widely separated than are any of the others, i.e. they 
are more laterally placed, so that in dorsal aspect even the endopodites are visible 
(PL 3 and Text-fig. 14) ; in ventral aspect they are concealed by the last pair of 
thoracic appendages. In other words, abdominal somites 2 to 5 are considerably deeper 
than somite 1 and their tergites bend in somewhat to meet their respective sternites, 
thus the pleopods are more ventral in position. A most unusual feature is the 
presence of a transverse lamella projecting freely backwards from the posterior 
sternal margin of each abdominal somite 3, 4 and 5. This chitinous duplicature, as 
Caroli terms it, becomes progressively longer (that on somite 3 being the shortest, 
that on 5 the longest). The protopodites of each pair of pleopods are incorporated 
with the respective lamella, but the muscle bands which work the two rami of each 
pleopod are visible through the chitin (Text-figs. 8, 14 and 15 m). Under certain 
lighting conditions, at a certain angle, there appears to be a median notch (an inverted 
V) dividing the lamella on somite 3 of the smaller paratype ; when a fine needle point 
is run along under the free margin of the lamella, however, the posterior margin is 
found to be continuous. 

One of the most distinctive features of this animal is the uropod (Text-figs. 8, 
12 and 13). The protopodite is expanded medially (e), beyond the base of the endo- 
podite (en), so that it occupies very nearly half of the posterior margin of the sixth 
abdominal somite (Text-fig. 8). Moreover, in ventral aspect, it has a long, massive, 



ON A STYGIOMYSIS FROM THE WEST INDIES 



295 




Figs. 12-14. Stygiomysis holthuisi (Gordon). Smaller paratype. Fig. 12. Left uropod, 
in dorsal aspect. Fig. 13. Same, in ventral aspect. The muscle bands within the proto- 
podite are restricted to the stippled area. Fig. 14. Abdomen and telson, in lateral aspect. 
e. Median expansion of protopodite. en. Endopodite. ex. Exopodite. m. Membranous 
lamellae connecting protopodites of pleopods 3, 4 and 5. p. Backward prolongation of 
protopodite. 1 and 6. First and sixth abdominal somites. 



296 



ON A STYGIOMYSIS FROM THE WEST INDIES 



backward prolongation (p), which extends almost to the posterior margin of the 
endopodite and bears five strong apical and subapical spines. These spines curve 
inwards and upwards (dorsally) towards those on the other partner. On the distal 
half of the inner margin of p there is a row of spinules interrupted by one or two 
spines. The left uropod of the smaller paratype was removed and is represented, 
in dorsal and ventral aspects, in "Text-figs. 12 and 13. In dorsal aspect only the proxi- 




Figs. 15-16. Stygiomysis holthuisi (Gordon). Fig. 15. Third left pleopod of female, and 
left half of membranous lamella, in ventral aspect. Fig. 16. Second pleopod of holotype 
(female), m. Lamella. 3. Side of third abdominal somite. Fig. 16 at larger scale. 



mal part of the protopodite with its medial expansion (e) is visible. The endopodite 
conceals part of the exopodite, and the long marginal plumose setae on exopodite 
and endopodite (mostly omitted in Text-fig. 12) conceal the apical spines on the 
prolongation p. There is no suture line or diaerisis on the exopodite, at the level of the 
spines on the outer margin, marking off the distal portion. The uropods are unusually 
stiff (at any rate in fixed specimens) ; when I tried to spread out the exopodite 



ON A STYGIOMYSIS FROM THE WEST INDIES 



!97 



from beneath the endopodite it was difficult to do so without damage to the exopodite 
base (Text-fig. 13 ex). There is no statocyst either in the uropod or in the basal 
segment of the antennular peduncle. 

Oostegites. In the holotype there are four pairs of developing oostegites arising 
from the coxae of thoracic limbs 4 to 7 respectively. The two anterior pairs are 
equally developed, each partner in the form of a long narrow lamella. The third 
pair is unequal, the left partner being appreciably larger than the right one ; of the 
last pair the left one is very small and the right one has not as yet appeared. 

Note on the male. On 27th October, 1958 I received from Dr. Holthuis another 
specimen which had been collected in the type locality on 25th May, 1958 by Mr. 
M. Vroman. This specimen, which proved to be a male, is rather more slender 
than the holotype and measures 10 mm. in body length. It is less contracted than 
the type specimens and not quite so flattened dorsoventrally (Text-fig. 24). The 
ratio of carapace length to total body length is perhaps more as in the living animal 
(1 : 474 in the male, as against 1 : 3-93 in the holotype). Apart from the more slender 
build, the presence of short penes, and the sexual dimorphism exhibited by the 
second pair of pleopods, I find no obvious differences between the male and the 
holotype — the antennula is often modified in male Mysidacea. Pleopod 2 is repre- 
sented in Text-fig. 18, that of the holotype in Text-fig. 16. As far as I can see without 
actually removing one of these pleopods from each specimen, the exopodite is three- 
segmented in the holotype and only two-segmented in the male. In the latter there 
are 8 or 9 long, strong, thorny setae (or spines) on the inner margin of the proximal 
segment and, in addition, 5 to 7 shorter setae which are more oblique (directed 
more ventrally) — see Text-fig. 18. The first pleopod is represented in Text-fig. 17. 

The genital opening is situated on a very low papilla at the base of the last thoracic 
appendage, on the inner side of the coxa. There is no coxal lamina such as Villalobos 
(1951, p. 208, Text-fig. 12) has figured for his genus Typhlolepidomysis. 

Remarks. These specimens from the West Indies are undoubtedly referable to 
Caroli's genus Stygiomysis, as a comparison of Text-fig. 8 with his Text-fig. 3 (1937, 
p. 223) shows. The type species of the genus, 5. hydruntina Caroli, was found in 
two grottos in the region of Otranto, Italy (the male holotype in the " Buco dei 
Diavoli ", one female and two immature specimens in " l'Abisso "). Caroli's brief 
description, based chiefly on the holotype, makes known the most important charac- 
ters of the species, and indeed of the genus. As far as I know, the more detailed study 
which was to have followed (Caroli, 1937, p. 220) has never been published. 

In addition to the photograph of the whole animal in lateral aspect, Caroli figured 
only the first gnathopod (second thoracic appendage) and the posterior end of the 
body in ventral aspect. Most of his description of S. hydruntina would apply equally 
well to 5. holthuisi. Nevertheless, the two species differ in certain respects. For 
example, in 5. holthuisi segment 5 of the endopodite of gnathopod 1 (that which 
bears the terminal claw) is much longer than in 5. hydruntina and the terminal claw 
and the pair of spines flanking it are very much shorter (in hydruntina they are much 
longer than the segment that bears them — cf. Text-fig. 9 with Caroli, 1937, fig. 2, p. 
222) . In S. holthuisi the terminal claw of peraeopod 4 (last thoracic limb) is rather shorter 
than segment 5, whereas in S. hydruntina the claw is longer than the dactylopodite. 

ZOOL. 6, 5 2o§§§ 



298 



ON A STYGIOMYSIS FROM THE WEST INDIES 



The shape of the uropod differs appreciably in the two species ; in S. hydruntina 
the outer margin of the prolongation p of the protopodite is distinctly concave, 
the exopodite is much broader, with a more semicircular apex (cf. Text -figs. 8, 12 
and 13 with Caroli, 1937, p. 223, fig. 3). 

According to Caroli, the pleopods of the male increase very slightly in length from 




Figs. 17-18. Stygiomysis holthuisi (Gordon). Male. Fig. 17. 

Second (modified) pleopod. 



First pleopod. Fig. i< 



the first to the fifth pair, but " nessuno di essi mostra particolari modificazioni 
sessuali ". In S. holthuisi, however, the second pair of pleopods in the male are modi- 
fied ; as far as I can ascertain without removing one of these pleopods, the exopodite 
has only two, instead of the usual three, segments, and the proximal segment has 
a row of 9 long spines as represented in Text-fig. 18 (compare with the first male 
pleopod, Text-fig. 17 and with the second female pleopod, Text-fig. 16). Also, the 
membranous flaps between the third, fourth and fifth pairs of pleopods differ in 






ON A STYGIOMYSIS FROM THE WEST INDIES 299 

the two species. Caroli says that, in each of these three pairs, " the peduncle (proto- 
podite) is completely fused with a wide duplicature of the integument, in the form 
of an almost semicircular lamina, movably articulated with the posterior margin 
of the sternite, [extending] from the external margin to the middle line, where it 
meets that of the other side " (my translation). He adds that the peduncle proper 
is indicated by the presence of the muscle bands which work the two rami, but that 
there is no suture between it and the duplicature (Caroli, 1937, pp. 223-224 and 
fig. 3). In S. holthiiisi these duplicatures unite to form a single membranous flap, or 
lamina in which the muscle bands are visible at either side (Text-figs. 8 and 15). 
I thought at first that there was a median slit separating the two halves of the mem- 
brane of the third pair of pleopods of the smaller paratype of S. holthiiisi. But, on 
re-examination, I find that there also the membrane is continuous although it is 
slightly creased in the middle (Text-fig. 14). 

The female specimen of S. hydruntina measured only 7 mm. in length, but it 
also had 4 pairs of developing oostegites arising from thoracic limbs 4-7. These 
oostegites were probably not quite so well developed as those in the holotype of 
5. holthiiisi (c .1. = 9 mm.). Caroli is almost certain that the full complement of 7 
pairs will be found in Stygiomysis. If not, then Stygiomysis will be quite exceptional. 
Seven pairs of oostegites are found in the Lophogastrida (Lophogastridae and Euco- 
piidae), in the Lepidopidae (all three genera), Petalophthalmidae and Boreomysinae. 
All the rest of the Mysida (Thalassomysis excepted) have only two or three pairs of 
oostegites arising from the last two or three pairs of thoracic appendages. It is to 
be hoped that an ovigerous female of either species of Stygiomysis will soon be found, 
so that the exact number of oostegites can be ascertained. 



II. ON THE SYSTEMATIC POSITION OF THE ST YGI OM YSI D AE 

Up to the present time the following cavernicolous Mysidacea have been described 
or mentioned in the literature : — 

Statocyst absent 

Lepidopidae : Lepidops servalus (Fage) 1924, 1925. Zanzibar. 

Spelaeomysis bottazzii Caroli 1924. Italy. (Found a second time 
by Stammer — see Caroli 1937, p. 226). 

Typhlolepidomysis quinterensis Villalobos 1951. Mexico. 
Stygiomysidae : 

Stygiomysis hydruntina Caroli, 1937. Italy. 

Stygiomysis holthiiisi (Gordon) 1958. West Indies. 

Statocyst present 

Mysidae : Heteromysis cotti Caiman 1932. Lanzarote. 

Troglomysis vjetrenicensis Stammer 1933, 1936. Herzegovina. 

Antromysis cenotensis Creaser 1936. Yucatan. 

Antromysis sp. Bolivar y Pieltain 1943-44, and 1950. Cuba. 



3 oo ON A STYGIOMYSIS FROM THE WEST INDIES 

Most of these species occur in fresh or brackish water ; Heteromysis cotti is a 
marine species since the cave in which it lives communicates with the sea. (Antro- 
mysis anophelinae W. M. Tattersall, 1951, p. 230, was discovered in the holes made 
by a land crab, Cardisoma crassum Smith, along with mosquito larvae, far up the 
mangrove inlets behind Puntarenas, Costa Rica — in water which must have been 
nearly, if not quite fresh) . The cavernicolous species with a statocyst in the endopo- 
dite of the uropod are referred to the large family Mysidae, in the suborder Mysida. 
Those that have no statocyst are peculiar or aberrant in certain respects and their 
systematic position is less easy to determine. Fage referred the family Lepidopidae 
to the suborder Mysida while admitting that it had a number of Lophogastrida 
characters ; Caroli thought the family might be regarded as modified or aberrant 
Lophogastrida and Villalobos thought that it should be placed in a new suborder 
(see p. 306). Caroli had no hestitation in referring the Stygiomysidae to the suborder 
Mysida. 

General appearance. In all three genera referred to the family Lepidopidae 
(formerly Lepidophthalmidae) the carapace agrees with the accepted definitions of 
the order Mysidacea. Caiman (1909, p. 171), in his definition of the order writes 
" the carapace extends over the greater part of the thoracic region, but does not 
coalesce with more than three thoracic somites ". Under morphology lower down 
on the same page he adds " from five to seven of the thoracic somites are distinct, 
and the last two or three may be left uncovered by the carapace on the dorsal side ". 
Caiman's definition is adopted by Tattersall, (1951, p. n) and that given by Zimmer 
(1927, p. 609) is in agreement with it. It has been altered by Dr. Olive Tattersall 
to read " carapace shield-like, covering the greater part of the thoracic region, 
fused dorsally with the head region and not more than four anterior somites including 
the first, which is incorporated in the head " (Tattersall and Tattersall, 1951, p. 11). 
Dr. Tattersall, in a letter dated 12th November, 1958, informs me that, as far as she 
can make out, " in some Mysidae there does appear to be fusion of at any rate part 
of the fourth somite with the carapace (in e.g. Mysidopsis, some Siriella and in 
Neomysis). I do not know quite how much one can rely on the " saddle " on the 
carapace in so many mysids, but I always considered it to represent the area of 
fusion of the two surfaces, i.e. the inner surface of the carapace with the dorsal 
surface of the thoracic somites involved." 

I have examined specimens of Lepidops servatus (Fage) but they are delicate 
and none too well preserved, and it is not easy to discover exactly how many thoracic 
tergites are free. As in most Mysidacea, the anterior thoracic somites are short and 
rather crowded together, especially dorsally, and the suture lines are visible laterally. 
As far as I can see four are certainly free and perhaps five. The last two tergites, 
which are uncovered by the emarginated carapace, are each as long as the first 
abdominal somite. The large lateral " wings " of the carapace extend to the first 
abdominal somite, and can easily be moved away from the body. All three genera 
retain the " caridoid facies " as do most Mysidacea. 

Stygiomysis, on the other hand, does not agree with Caiman's definition, even as 
emended by Dr. Tattersall. The carapace is unmistakably fused with the first 
four thoracic tergites but has no " wings " to speak of, so that the last four thoracic 



ON A STYGIOMYSIS FROM THE WEST INDIES 301 

somites are entirety exposed (PL 3, Text-figs. 1 and 24). Moreover, the body behind 
the short carapace (the sides of which cannot easily be separated from the body — at 
any rate in preserved specimens) is vermiform and very flexible. Caroli's photograph 
of Stygiomysis hydruntina shows the abdomen curved gracefully upwards, in a most 
un-mysid-like position (1937, p. 221, fig. 1). As that author says, the short carapace 
and the apparent division of the thorax into two subtagmata gives the genus "un 
aspetto particolare ". The sixth abdominal somite is distinctly shorter than either 
the fifth or the fourth somite. In Mysidacea " the sixth abdominal somite is usually 
longer than any of the others " (Tattersall and Tattersall, 1951, p. 16). This is the 
chief reason why I at first thought that the genus could not be referred to the order 
Mysidacea as at present defined (Gordon, 1958). 

The other two most striking peculiarities of the genus Stygiomysis namely, the 
transverse membranes with which are incorporated the protopodites of pleopods 
3, 4 and 5, and the long backward extension of the protopodite of the uropod, 
are not quite as unique as I had supposed. Each will be dealt with in turn. 

Transverse chitinous lamellae. Caroli (1937, p. 225) found what he called 
homologous though not identical structures in Lepidops and Spelaeomysis. I have 
been able to confirm his observation in Lepidops ; between the bases of pleopods 3, 
4 and 5 there is a transverse laminar extension from the posterior margin of each 
abdominal somite, as represented in Text-fig. 20. These chitinous duplicatures of 
the integument, as Caroli calls them, increase in length from somites 3 to 5 but the 
protopodites of the pleopods are not incorporated with them. In Spelaeomysis 
they are said to be even better developed, with a median emargination in the posterior 
border. Caroli assumes from this that each lamina had a double origin— i.e. that it 
arose as two duplicatures of the integument which had fused medially (Caroli, 
1937, p. 225, cf. also his fig. 3). Unfortunately, Villalobos (1951) does not state 
whether or not these laminae are present in Typhlolepidomysis ; they were overlooked 
when Lepidops and Spelaeomysis were first examined and could easily have been 
overlooked a third time. (But see footnote to p. 308). 

Although transverse laminae are known only in these more or less aberrant 
Mysidacea among the Malacostraca, they recall somewhat analogous structures in 
the Copepoda. Here it is the thoracic limbs which are involved ; " the proximal 
segments of the protopodites of each pair are connected with each other across 
the middle line by a plate formed by a transverse fold of the sternal integument . . . 
so that in the backward and forward movement in swimming the two appendages 
move as one " (Caiman, 1909, p. 81). See, for example, the figures of the reduced 
last limb of male Harpacticids (P5 <J) in Chappuis 1931, p. 521, fig. 13 ; p. 538, 
fig. 60 — these limbs are not always joined in the female. 

How do the duplicatures arise in the Mysidacea? In Lepidops the chitinous 
lamina seems to be simply a backward extension or fold of the posterior sternal 
margin of the abdominal somite, since the protopodites of the pleopods are not 
involved (Text-fig. 20, m). In Stygiomysis hydruntina, Caroli's figure (1937, fig. 3) 
suggests that each lamina may have a double origin — that each half arises as a 
median expansion of the protopodite of the pleopod which fuses anteriorly with the 
sternal margin of the abdominal somite, since the two halves merely meet in the 



302 



ON A STYGIOMYSIS FROM THE WEST INDIES 




ON A STYGIOMYSIS FROM THE WEST INDIES 303 

middle line. In Stygiomysis holthuisi the lamina might arise in either of these ways, 
since the protopodites of the pleopods are incorporated and there is no trace of 
a median suture line (see also p. 294). It would be interesting to know how these 
chitinous laminae develop and what their function can be, since the pleopods are 
almost certainly not natatory. The very thin sternal wall above each lamella may 
be respiratory, 

Uropod. In Lepidops servatus the protopodite is broad, occupying half of the 
posterior margin of the last abdominal somite. Moreover, on the ventral side, there 
is a short but distinct backward prolongation armed with a terminal spine and with 
numerous spinules on the inner border (Text-fig. 21, p). This might represent an 
initial stage in the development of a massive backward prolongation such as is 
found in Stygiomysis, although Caroli prefers to regard it as a last vestige retained 
in the family Lepidopidae (1937, p. 225). At any rate, this incipient or vestigial 
structure in Lepidops (and presumably also present in Spelaeomysis), together with 
the transverse laminae mentioned above, suggest a fairly close relationship between 
the Lepidopidae and the Stygiomysidae (see also p. 308). A somewhat analogous 
structure is characteristic of the uropod of the Stomatopoda, which might indicate 
that the prolongation of the protopodite is archaic and, as Caroli thinks, may at 
one time have been much more widely distributed among Malacostraca. I am 
inclined to think that it is a highly specialized structure which has arisen indepen- 
dently in the Stygiomysidae, and to a lesser extent in the Lepidopidae, in response 
to some special habit. 

It may be mentioned in passing that, in the genus Micraspides (Syncarida, Anaspi- 
dacea), the uropod is also armed with strong spines, not on the protopodite (which is 
not prolonged) but on the endopodite. On the dorsal surface, near the postero- 
median angle, are three strong spines directed obliquely upwards ; in addition there 
is a row of special spines on the inner margin (Text-fig. 19, en). Nicholls' figure 
(193 1, pi. 32, fig. 15) of the uropod of the type species Micraspides calmani is rather 
small and does not show the row of spines near the external margin of the exopodite. 
In Thermosbaena mirabilis Monod the exopodite of the uropod is remarkably spiny 
(Monod, 1940, fig. 19). 

Pleopods. The pleopods of Stygiomysis, though smaller and not increasing 
much in length from the first to the fifth pair, are of the same general form as those of 
Lepidopidae (PL 3, and Text-fig. 14, cf . Text-fig. 20) . In both families they are reduced, 
non-natatory, and biramous in both sexes. In Stygiomysis holthuisi the second pair 
is modified in the male although it is not longer than the others (Text-figs. 17 and 
18) ; Caroli's statement that there is no sexual dimorphism of the pleopods in 



Figs. 19-21. Micraspides calmani Nicholls. Fig. 19. Left uropod of syntype, in dorsal 
aspect, omitting long marginal setae. Lepidops servatus (Fage). Syntype from Paris 
Museum. Fig. 20. Abdominal somites 3 to 6, and protopodites of uropods, in ventral 
aspect, showing membranous free laminae (m) on posterior margin of somite 3, 4 and 5, 
between bases of the pleopods. Fig. 21. Left uropod, in ventral aspect (endopodite 
damaged), en. endopodite. ex. Exopodite. p. Short backward prolongation of proto- 
podite. Scale = 05 mm. 



3 o 4 ON A STYGIOMYSIS FROM THE WEST INDIES 

5. hydruntina requires corroboration. The second pleopod of the male is similarly 
modified in all three genera of the Lepidopidae — see Fage, 1925, fig. VI, 2 and 2 J; 
Caroli 1924, p. 513 ; Villalobos 1951, fig. 13, A and B. (A glance at fig. 1 of Villalobos' 
paper shows that pleopod 2 is shorter than pleopod 5 in the male holotype ; it 
seems, therefore, that in his fig. 13, C and D are drawn at a smaller scale than A 
and B). Pseudobranchiae are absent in both Stygiomysidae and Lepidopidae. 

In Lophogastrida the pleopods of both sexes are well developed, biramous, multi- 
segmented and natatory ; they may be considerably larger in the male than in the 
female, but are unmodified or very slightly so. In Mysida they are seldom alike in 
both sexes and generally exhibit marked sexual dimorphism. In the female they 
are, as a rule, rudimentary, uniramous (rarely the first may be biramous) and non- 
natatory ; occasionally those of the male are all rudimentary as in the female 
(Mysidella, Mysidetes, Heteromysis). In the male they are sometimes biramous, 
natatory and unmodified (Petalophthalmidae) , or the third, or second and third, 
pairs are slightly modified (Boreomysinae) ; more often one or more pairs are reduced 
as in the female and one or more pairs are modified as accessory copulatory organs ; 
pseudobranchiae are present. The pleopods which are most commonly modified are 
the third and/or fourth pairs. In the genus Rhopalophthalmus it is the second pair 
(Tattersall, O.S. 1957, fig. 7H, compare with G, J, K and L) ; but here the pleopods 
are biramous, multiarticulate, with conspicuous pseudobranchial lobes at the bases 
of the endopodites. The modified pair, or pairs, of male pleopods are usually long 
and conspicuous in the family Mysidae, to which Rhopalophthalmus belongs. It is 
just a coincidence that the same pair of male pleopods is modified in this genus as 
in the cavernicolous families under discussion. 

Mouthparts. There are numerous differences in detail between the individual 
mouthparts of the two families and these are doubtless correlated with differences 
in feeding habits. For example, in the Lepidopidae the molar process of the mandible 
is better developed, the palp is larger and the second and third segments are each 
armed with a close row of thorny spines (cf. Text-fig. 27, where many of the spines 
and setae are omitted, Fage, 1925, fig. IV, M and Villalobos, 1951, fig. 6 with figs. 4 
and 5). 

According to Tattersall and Tattersall, (1951, pp. 20-21), in Mysidacea the maxil- 
lula is smaller than the maxilla and always lacks a backwardly directed palp or 
endopodite except in the genus Gnathophausia. This statement requires amplifica- 
tion however, because there is a palp in all three genera of the Lepidopidae (Fage, 
1925, fig. IV, m 1 ; Caroli, 1924, p. 513 and Villalobos, 1951, fig. 7). In each of three 
specimens of Lepidops servatus which I have examined, this palp has a median 
constriction and appears, therefore, to be two-segmented as in Typhlolepidomysis 
(see Text-figs. 27 and 28). Moreover, in Stygiomysis holthuisi the maxillula is larger 
than the maxilla ; the outer endite is stout and one of its apical spines is greatly 
enlarged ; there is no palp (Text-figs. 2 and 5 ; cf . the small maxillula of Lepidops 
represented in Text-figs. 28 and 27). 

The maxilla of Stygiomysis has strong spines rather than setae on the whole inner 
margin but they do not continue along the lower border of the simple endite (Text- 
fig. 6, on en, d and s) ; in the Lepidopidae the inner and lower margins of the maxilla 



ON A STYGIOMYSIS FROM THE WEST INDIES 305 

bear numerous finer setae and spines, and the segments of the endopodite are much 
longer (Fage, 1925, fig. IV, m 2 and Villalobos, 1951, fig. 8). The first thoracic append- 
age or maxilliped of Stygiomysis is more pediform, lacks the large endites, but has 
strong spines on the inner margin of the distal segments of the endopodite (Text-fig. 
7 ; cf. Fage, 1925, fig. VI, mp and Villalobos, 1951, fig. 9). According to Siewing 
(1958, pp. 250-251) this appendage should be called a gnathopod in the Stygio- 
mysidae and a maxilliped in the family Lepidopidae (see later, under terminology, 
p. 316). Differences such as these may be found between closely related families 
of Mysidacea. The exopodite of the first thoracic limb is reduced to a simple unseg- 
mented lamina, which is relatively smaller in Stygiomysis holthuisi than in either 
Lepidops or Typhlolepidomysis (cf. Text-fig. 7 with Fage, 1925, fig. VI, mp and Villa- 
lobos, 1951, fig. 9). In this respect both families are nearer to the suborder Lopho- 
gastrida and to the Petalophthalmidae (in which the exopodite is lacking, as in some 
species of Gnathophausia) than to the Mysidae. 

Labium. The paragnatha in Stygiomysis are unusual and very conspicuous, 
although Caroli did not mention them. To discuss the labium at this point would 
necessitate rather a lengthy digression ; it will be dealt with in a separate section 
later on (p. 309). Of the paragnatha that I have examined, those of Lepidops seem 
to be nearest to those of Stygiomysis. 

Thoracic appendages 2-4. Many of the peculiarities of the mouthparts men- 
tioned above are doubtless correlated with the fact that in Stygiomysidae three 
pairs of thoracic appendages (2-4) are modified as gnathopods, whereas in Lepi- 
dopidae only one pair, the second, is thus modified (cf . PL 3, Text-figs. 9 and 10 with 
Fage 1925, fig. V, p 2 and Villalobos, 1951, fig. 10). Incidentally, Villalobos refers to 
the second thoracic limb as " primer par de pereiopodos " in the text and as " maxil- 
lipedo del secundo par " in the legend to fig. 10 and Caroli (1937, p. 226) says 
" toracopodi 2-4 (gnathopodi) massillipediformi "! Only in the family Eucopiidae 
of the suborder Lophogastrida are the same three pairs of thoracic limbs modified 
as gnathopods ; but in this family the following three pairs of appendages are 
elongated and specially modified (Tattersall and Tattersall, 1951, p. 111, figs. 13C, 
D and E and 12A). When Sars (1885, p. 59) suggested that the long prehensile 
thoracic limbs 5 to 7 might be used for seizing hold of Hydroids or Crinoids fixed at 
the sea bottom, he was unaware of the fact that the Eucopias are bathypelagic 
(Fage, 1 941, p. 3 and 1942 pp. 3 and 60). There is nothing in mid- water for a bathy- 
pelagic animal to cling to ; these prehensile endopodites, reaching well beyond the 
body, seem admirably suited for keeping the animal suspended and for seizing prey 
which could be passed to the short robust endopodites of the gnathopods (see p. 319). 
Stygiomysis is probably raptorial in its feeding habits whereas the Lepidopidae are 
probably predominantly filter feeders. 

Oostegites. In Lophogastrida, Lepidopidae, Petalophthalmidae and Boreomy- 
sinae (Mysidae) a pair of oostegites or brood lamellae arises from each pair of thoracic 
appendages 2-8. Throughout the rest of the large family Mysidae there are either 
three or two pairs, arising from the last three or two pairs of thoracic appendages. 
The only exception to this rule is the genus Thalassomysis Tattersall 1939, p. 238 
(established two years after Caroli's genus Stygiomysis). Here reduction has not 



3 o6 ON A STYGIOMY SIS FROM THE WEST INDIES 

gone quite as far and four pairs of oostegites are present. Of these the first pair is 
vestigial, the second pair small and the last two pairs form the major part of the 
brood chamber. Oostegites disappear from the more anterior limbs first so that, 
no matter what their number, a pair is always present on the seventh and the eighth 
thoracic appendages. In Stygiomysis the full complement is unknown but, unless 
the genus is quite exceptional, there ought to be a pair on the eighth thoracic limbs ; 
this means that there ought to be at least five pairs and Caroli may be right in 
thinking that there may be seven pairs. 

Discussion. The family Stygiomysidae seems to be distinct from, but most 
closely related to, the family Lepidopidae. But, as already mentioned, opinion 
differs as to the systematic position of the family Lepidopidae. Since gills are 
absent and the pleopods are reduced, non-natatory, Fage (1924, 1925) referred his 
new family " Lepidophthalmidae " to the sub-order Mysida (Mysidea of some 
authors), at the same time stressing its affinities with the sub-order Lophogastrida. 
Both Caroli and Villalobos have raised objections to this. Caroli's announcement 
of Spelaeomysis appeared before Fage's main paper (in December, 1924). He says 
that the discovery of a second genus in Italy, so far from the habitat of the first 
genus (Zanzibar), shows that these forms must have had at one time a very wide 
distribution. " Per 6 e forse prematuro ritenerle come forme di passagio dai Lopho- 
gastrida ai Mysida. A mio avviso, potrebbero anche essere considerate Lophogastrida 
modificati (perdite delle branchie, riduzione dei pleopodi) dalla vita cavernicola 
assunta in epocha remota " (Caroli, 1924, p. 513). That Fage adheres to his original 
opinion is evident from a remark he makes in a later work on Lophogastrida (1941, 
p. 3) ; discussing some primitive characters he mentions " le palpe retro verse, 
herite des Cumaces et des Tanaidaces, que portent au premieres maxillae les Gnatho- 
phausia. Seuls parmi les Mysides [Mysida] les genres Lepidops (Lepidophthalmus 
Fage 1924) et Spelaeomysis Caroli 1924 en sout pourvus, qui revelent ainsi leur part 
du meme heritage ". Villalobos (1951, p. 216), after a brief discussion of the syste- 
matic position of the Lepidopidae concludes : " Segun nuestro punto de vista, la 
position adoptada por Fage es vm poco forzada, porque si los caracteres distinctivos 
de la familia se encuentran representados ya sea en Lophogastridea o en Mysidea, 
lo convenient e seria crear un soborden intermedio ". He did not, however, propose 
a new sub-order because of the paucity of material and the small number of genera 
and species known (three monotypic genera). 

When Caroli (1937, p. 225) discovered the transverse lamellae and a short but 
distinct backward prolongation of the protopodite of the uropod in Lepidops and 
Spelaeomysis (see Text-fig. 20, m and Text-fig. 21, p of this paper) he said that he 
could not judge whether the presence of these two characters indicated a closer 
affinity between the Lepidopidae and Stygiomysis. " Si puo anche suppore che si 
trata di caratteri arcaica, communi probabilmente in passato ad altre e piu numerose 
forme, e conservatisti, benche in via di graduale riduzione, nei nostri Misidacei, 
in grazzia dello speciale habitat ". He does not explicitly say that he has altered his 
earlier opinion (1924) namely, that the Lepidopidae are perhaps modified Lopho- 
gastrida, and he has no hesitation in referring the Stygiomysidae to the sub-order 
Mysida. 



ON A STYGIOMYSIS FROM THE WEST INDIES 



307 



The family Lepidopidae shares its diagnostic characters with the rest of the Mysi- 
dacea as follows : — 

(i) Gills absent : Stygiomysidae and the whole of the Mysida. 

(ii) Oostegites seven pairs : Lophogastrida, Petalophthalmidae, Boreomy- 
sinae and ? Stygiomysidae. 

(iii) Statocyst absent : Lophogastrida, Stygiomysidae and Petalophthalmidae. 

(iv) Palp present on maxillula : Gnathophausia only (Lophogastrida). 

(v) Exopodite of maxilliped (first thoracic limb) a small unsegmented lamella 
or absent : Lophogastrida, Stygiomysidae and Petalophthalmidae. 

(vi) Second thoracic appendage modified as a gnathopod : most Lopho- 
gastrida and Mysida — in the Stygiomysidae and Eucopiidae the next two pairs 
are similarly modified. 

(vii) Endopodites of thoracic appendages 3-8 (or 5-8) without subdivision 
of the penultimate segments : Lophogastrida, Stygiomysidae, Petalophthal- 
midae and a few Mysidae. 

(viii) Pleopods reduced, non-natatory, but biramous in both sexes ; second 
pair slightly modified in (J ; Stygiomysidae (see p. 303). 

(ix) Transverse lamellae arising from the posterior sternal margins of 
abdominal somites 3, 4 and 5 ; Stygiomysidae, though here the protopodites 
of the pleopods are also incorporated in the lamellae. (Text-figs. 20 and 8 and 
Caroli, 1937, fig. 3). 

(x) A short but distinct backward prolongation of the protopodite of the 
uropod : Stygiomysidae, here it is very much longer (Text-figs. 21 and 13, p). 

(xi) Exopodite of second thoracic limb normal, multiarticulate : most 
Mysidacea (except Petalophthalnuis , where it is absent). It is long and natatory 
as in forms with reduced pleopods. The basal segment of this and the following 
exopodites are long and narrow as in Stygiomysis and Lophogastrida, not 
blade-like as in most Mysida. 

Certain characters mentioned by Fage in his diagnosis of the family Lepidopidae 
are of less importance in trying to elucidate affinities since they may vary within 
one and the same family : (a) Endites or lobes on the first thoracic endopodite may 
be large, poorly developed or absent, depending on the feeding habits. They are 
well developed in all three genera of Lepidopidae, absent in Stygiomysidae ; in 
the family Mysidae they are large in Mysis, one only is present in Mysidetes and they 
are absent in Mysidopsis. In the Petalophthalmidae they are absent in Hansenomysis, 
present in Petalophthalmus. In the Lophogastridae they are present in Gnathophausia 
(Text-figs. 33-34), absent in Lophogaster. (b) Presence or absence of a suture line 
(diaerisis) on the exopodite of the uropod. In the Lepidopidae there is one in Lepidops 
(which only shows up well in reflected light) ; it is said to be present in Typ/ilo- 
lepidontysis, absent in Spelaeomysis. It is present in the majority of the Lopho- 
gastrida and in Petalophthalmidae, is faint and incomplete in Siriellinae, absent 
in other subfamilies of Mysidae and in Stygiomysidae. Rholapophthalmus is excep- 
tional in having a suture in both exopodite and endopodite. 



3 o8 ON A STYGIOMYSIS FROM THE WEST INDIES 

From the above general survey of the characters used to subdivide the order 
Mysidacea, it is obvious that, as far as external morphology is concerned, the only 
constant character separating the Lophogastrida from the Mysida, Lepidopidae 
and Stygiomysidae, is the presence of podobranchiae or gills on some or all of the 
thoracic limbs. A negative character like the absence of gills does not necessarily 
imply that the Lepidopidae and the Stygiomysidae belong to the Mysida — as Caroli 
1924, p. 513 implies, loss of gills may have occurred more than once in the course of 
evolution. Several of the Lophogastrid characters are shared by the Lepidopidae, 
Stygiomysidae and Petalophthalmidae and the large number of oostegites is shared 
by the Boreomysinae also (as are natatory pleopods in the male). But I do not see 
that anything is to be gained by erecting a third suborder for the Lepidopidae 
and Stygiomysidae. In that case something would have to be done about the Petal- 
ophthalmidae and perhaps the Boreomysinae as well. In the meantime, I prefer 
to leave both families (Lepidopidae and Stygiomysidae) in the Mysida since podo- 
branchiae are absent and the pleopods are reduced. These two families are very 
closely related to each other and differ in several important characters from the rest 
of the Mysida — the similar pleopods, the transverse lamellae on abdominal somites 
3, 4 and 5 and the greater or lesser prolongation of the protopodite of the uropod. 
Their relationship to each is much the same as that of the Lophogastridae and 
Eucopiidae. Whatever future specialists may decide to do with the Lepidopidae, 
I am convinced that the Stygiomysidae should be ranked along with it — even if 
there should be fewer than seven pairs of oostegites in Stygiomysis. 

The family Lepidopidae requires redefinition, because certain characters were 
overlooked at first ; also for comparison with that of the Stygiomysidae, which is 
the same as for the genus Stygiomysis (see p. 286). 

Diagnosis of the family Lepidopidae : Cavernicolous Mysidacea exhibiting 
the " caridoid facies". Carapace relatively long, smooth or with a sinuous furrow ; 
produced antero-laterally beyond the rounded rostral margin, postero-lateral 
" wings " well developed, reaching the first abdominal somite laterally, but leaving 
the last two thoracic tergites exposed dorsally. (A scale-like process on penultimate 
thoracic tergite in Typhlolepidomysis) 1 . Abdomen, including telson, half as long 
again as the cephalothorax, not broader in middle, sixth somite as long as, or longer 
than, fifth. Telson truncate or tapering ; spinules or spines on distal half of each 
lateral margin as well as on the rounded apex (spinulation not mentioned in Spelaeo- 
mysis). Ocular scales quadrangular, distal margin oblique, without or with traces 
of visual elements (a few ommatidia in Lepidops). Antennular and antennal flagella 
as long as (Lepidops) , or far exceeding, the length of the body (Typhlolepidomysis) 1 ; 
antennal scale well developed, with setae on inner and outer margins. Labrum 
oval, longer than wide (Lepidops). Labium with paragnatha well separated proxi- 
mally, but not as long as in Stygiomysidae. Mandible with well developed incisor 
and molar processes ; a row of strong thorny spines on second and third segments of 
the palp. Maxillula small, with a 2-segmented reflexed palp ending in a long curved 

1 Since this paper went to press I have collected specimens of Spelaeomysis bottazzii from the type 
locality. It has enormously long antennae and a scale behind the dorsal posterior emargination of the 
carapace. Typhlolepidomysis is therefore a synonym of Spelaeomysis. 24.xi.59. 



ON A STYGIOMYSIS FROM THE WEST INDIES 309 

seta ; all apical spines on the distal endite of about same size. Maxilla with setae, 
some of which are distally plumose ; segments of endopodite quite long. Maxilliped 
not pediform, with well developed endites and the penultimate segment broadened ; 
exopodite a small unsegmented lamella with or without fine marginal setules ; 
epipodite rather long and narrow. Second thoracic appendage with exopodite 
longer than endopodite, which may be modified as a gnathopod. Thoracic endopo- 
dites 3 to 8 all slender ambulatories. Penultimate segments of thoracic endopodites 
not subdivided. Pleopods alike in both sexes, apart from the slight sexual dimorphism 
exhibited by the second pair ; they increase in size from the first to the fifth pair, 
are biramous, with one-segmented endopodite and three-segmented exopodite 
(four-segmented in the second pair in the female). A short but distinct transverse 
chitinous lamella projects from the posterior sternal margin of abdominal somites 
3, 4 and 5, with which the protopodites of the pleopods are not incorporated. Uropod 
with a small but distinct backward prolongation of the protopodite, armed with a 
terminal spine (? overlooked in Typhlolepidomysis) ; statocyst absent ; a sinuous 
suture line on the exopodite (absent in Spelaeomysis). Seven pairs of oostegites. 

Type genus : Lepidops (Fage) 1924, 1925. L. servatus (Fage) is stated to be 
planktonic : " il existait dans ce petit lac de Machumwi-Ndogo un veritable plancton, 
compose de petits Schizopodes a yeux tres reduits. lis ne se sont pas fait prendre 
aux nasses, mais nous en avons receuilli un grand nombre pres de la surface, au 
filet fin ". Allaud et Jeannel in Jeannel and Racovitza (1914, p. 381). 



III. NOTE ON THE PARAGNATHA IN SOME MYSIDACEA 

As a rule, little or no attention is given to the labium or lower lip when the mouth- 
parts of Malacostraca are examined — unless, of course, the feeding habits are being 
studied in detail. It is scarcely surprising, therefore, that in the subject index of the 
Ray Society Monograph of the British Mysidacea (Tattersall & Tattersall, 195 1) 
the labrum or upper lip is included, but the labium is not mentioned under any of 
its synonyms (hypostoma, metastoma, paragnatha). In Stygiomysis, however, 
the paragnatha are so conspicuous that they could not possibly be overlooked 
(Text-figs. 1 and 3, / and Text-fig. 23, p) ; yet Caroli does not mention them in his 
description of S. hydruntina (1937). 

Michael Sars (1862, pi. 1, figs. 25 and 26) gave two minute sketches of the labium 
of Lophogaster typicus and G. O. Sars not infrequently figured the posterior, as well 
as the anterior lip. In his Challenger Report for example, he included small figures 
of the labium of Gnathophausia and Eucopia (Lophogastrida) , of Anchialus and 
Euchaetomera (Mysida) (Sars, 1885, pi. 8, fig. 5 ; pi. 9, fig. 8 ; pi. 34, fig. n ; pi. 35, 
fig. 7 and pi. 37, fig. 5). These suggest that the labium is more deeply cleft in the 
Mysida than in Lophogastrida. The paragnatha are also figured in situ for Hemimysis 
by Cannon and Manton (1927, pi. ill, fig. 4) and for Lophogaster by Manton (1928, 
pi. in, fig. 6) ; and there are others. But I do not recall having seen any illustration 
of the paragnatha quite like those of Stygiomysis. In lateral aspect, they are unusu- 
ally pendant, falling well below the level of the labrum, and the distal part of each 



3io 



ON A STYGIOMYSIS FROM THE WEST INDIES 




mm 




ON A STYGIOMYSIS FROM THE WEST INDIES 311 

is spoon-shaped. (Text-fig. 1, /). In ventral aspect, the paragnatha are widely 
separated proximally, and the spoon-shaped parts conceal much of the mandibles 
and may over-reach the posterior rim of the labrum (Text-fig. 3, / and Text-fig. 
23, p). I am not sure whether there is a subsidiary basal lobule or whether it is just 
a twist to form a sort of neck (n) or stalk for the spoon-like head. I have examined 
the paragnatha in several genera of Mysidacea for comparison with those of Stygio- 
mysis. But there is far more variation than I had anticipated and time is lacking 
for a detailed comparative study. Nouvel (1943) made a very careful study of the 
labrum in most of the species that he described and figured in his excellent Report. 
But, as a rule, he does not mention the labium except in Thalassomysis tatter salli 
in which he says that it is " tres aberrant. II est aussi tres proeminent et asymetrique. 
Sa par tie droit e distale [right paragnath] comporte une cavite ou vient se loger la 
dent du labrum. Sa partie gauche epouse aussi le contour du bord posterieur gauche 
du labrum. Presque toute la partie distale est couverte de petites soies analogues 
a celles du bord gauche du labrum " (p. 59 ; pi. in, fig. 89). But he does not say 
what he considers the most typical form, or forms, of the labium in the Mysidacea. 

In Eucopia sculpticauda Faxon (Eucopiidae-Lophogastrida) the labium, repre- 
sented in Text-fig. 32, is not quite as deeply cleft as that of Lophogaster (Lophogastri- 
dae) as figured by M. Sars (1862, pi. 1, fig. 25) ; in the micro-preparation the parag- 
natha are slightly displaced, owing to pressure of the cover slip, so that a secondary 
lobule, armed with marginal spinules, is visible (cf. Lophogaster in Manton, 1928, 
pi. in, fig. 6). 

Of the highly specialized yet primitive family Petalophthalmidae (Mysida) I 
have examined Petalophthalmus armiger W. Suhm and Hansenomysis fyllae (Hansen). 
In Petalophthalmus the labium is deeply cleft (Text-fig. 26, right-hand drawing) ; 
in ventral aspect, the paragnatha are much longer than wide, are separated proxi- 
mally by a narrow gap and meet distally. In lateral aspect, each paragnath is deeply 
concave, almost bowl-shaped, and the cutting edge of the mandible lies in the con- 
cavity (Text-fig. 26, left sketch, / and m) . In Hansenomysis the paragnatha are also 
longer than broad and in situ are not widely separated proximally (Text-fig. 29, 
upper sketch, made before mounting on a micro-slide). In the micro-preparation 
the paragnatha are displaced owing to pressure of the cover-slip and the inner lobules 
with their marginal spinules are visible (cf. Text-figs. 29 and 32). 

In Text-fig. 22 the labium of Boreomysis rostrata Illig (Boreomysinae-Mysida) 
is represented in three different aspects ; the paragnatha are slightly asymmetrical 
and there is no distinct subsidiary lobule on either of them. An inner knob or boss (a) 
can be seen in situ between the paragnatha (upper right sketch which is the aspect 



Figs. 22-24. Boreomysis rostrata Illig. Fig. 22. Labium or lower lip, in three different 
aspects. Stygiomysis holthuisi (Gordon). Fig. 23. Right half of labium, in ventral aspect 
(incisor edge of left mandible also shown) . Fig. 24. Anterior half of male, in dorsal aspect, 
(first abdominal somite represented by broken line), a. Boss on labium, n. Neck. 
p. Paragnath. 5 and 8. Fifth and eighth thoracic somites. 



312 ON A STYGIOMYSIS FROM THE WEST INDIES 

i i 



0.5 




Figs. 25-28. Petalophthalmus armiger W. Suhm. (Reg. no. i940,viii.5. 568-569). Fig. 
25. Right maxillula. Fig. 26. Labrum, right mandible, paragnath and maxillula, in 
lateral aspect; also right paragnath in ventral aspect. Lepidops servatus (Fage). 
Syntype from Paris Museum. Fig. 27. Labrum, labium, mandibles and maxillulae, 
in ventral aspect. Fig. 28. Same, in lateral aspect, at higher magnification. /. Parag- 
nath of labium, m. Mandible, mx. Maxillula. p. Mandibular palp. u. Labrum or 
upper lip. 



ON A STYGIOMYSIS FROM THE WEST INDIES 313 

from which most other paragnatha are figured). In the micro-preparation of Indo- 
mysis annandalei Tattersall (Mysini-Mysida) the labrum and labium are somewhat 
flattened by the cover-slip (Text-fig. 30, upper sketch) ; the labium is more deeply 
cleft than in Eacopia (Text-fig. 32) ; each paragnath is almost as wide as long and 
may touch the posterior margin of the labrum ; there is apparently no secondary 
lobule (upper and lower sketches). 

Of Lepidops servatus (Fage), which is a very minute and fragile species, I have 
examined a damaged specimen in which the paragnatha are flexed away from the 
mandibles and the right maxillula is somewhat displaced. The camera lucida sketches 
were prepared before any attempt was made to mount the fragment (Text-figs. 
27 and 28). The paragnatha are obviously rather widely separated proximally and, 
of all those examined, they approach most nearly to those of Stygiomysis although 
they are smaller and have no " neck " portion (or subsidiary lobule — see p. 311). 
In my preliminary note on " Rhopalonurus " holthuisi I referred to the paragnath 
or main lobe as a lappet (Gordon, 1958, p. 152) but it probably is not comparable 
with the small moveably articulated lappets found in some Tanaidacea. The proxi- 
mal portion (Text-fig. 23, n) may represent the subsidiary lobule which is present 
in Eucopia and in Hansenomysis (Text-figs. 32 and 29 — in each the lobule is beset with 
spinules) — or it may just be the result of a slight twist to form a " neck " or stalk. The 
unusual form and size of the paragnatha in Stygiomysis is probably a secondary 
specialization connected with its feeding habits, as are the other peculiarities of the 
mouthparts. The large maxillula with the remarkably enlarged spine on the distal 
endite has already been referred to ; I have figured the maxillula of most of the 
species of Mysidacea examined for comparison with that of Stygiomysis (cf. Text- 
figs 25-28 and 29, 30, 32 with Text-figs. 3 and 5). When the anterior (uppermost) 
spine on the distal endite is larger than the others this difference in size is not 
very pronounced (Text-figs. 25, 32 and 30). 

The feeding habits have been studied in detail only in a few species of Mysidacea 
although they have been deduced or inferred from an examination of the mouthparts 
in other species. But the significance of the marked differences in both form and size 
of the paragnatha, for example, is not known. A summary of our knowledge of 
food and feeding habits is given in Tattersall & Tattersall, 1951, pp. 29-33 and other 
references are listed in their subject index under feeding habits, food, filter feeding 
and raptorial feeding. Many Mysids employ the filtratory method and the raptatory 
method quite independently, e.g. Hemimysis (Cannon and Manton, 1927) and Gnatho- 
phausia, although in the latter there can be no auxilliary food current created by 
the thoracic exopodites as in other primitive filtratory Malacostraca (see Manton, 
1928, p. 117). Others are incapable of filter feeding and depend entirely on large 
food masses, like Lophogaster (Manton, 1928, p. no) and Petalophthalmus (Tattersall 
& Tattersall, 195 1, p. 113). The number of thoracic limbs adapted to aid in feeding 
varies considerably. In the Lepidopidae, as in many Mysids, the first and second 
pairs are modified as maxilliped and gnathopod respectively. But others may 
also assist, though not specially modified — e.g. in many Mysidae all the thoracic 
endopodites may assist in dealing with large food masses, although this is not their 
main function (Tattersall & Tattersall, 1951, p. 30). In Stygiomysis the first four 



314 



ON A STYGIOMYSIS FROM THE WEST INDIES 




0.5 



5 



Figs. 29-32. Hansenomysis fyllae (Hansen). (Reg. no. 1909.xii.24. 100-103) Fig. 29. 
Labium and maxillula, in ventral aspect (paragnatha displaced by pressure of cover slip) ; 
above, paragnatha figured before mounting the preparation. 

Indomysis annandalei Tattersall. (Reg. no. 1946. xi. 26. 45-64). Fig. 30. Labrum, 
labium, left mandible and maxillula ; below, right paragnath. (Preparation slightly 
flattened by cover slip) . Eucopia sculpticauda Faxon. (Reg. no. 1949. vii. 13. 65-75). 
Fig. 31. Right maxilliped, inner (or dorsal) surface. Fig. 32. Labium and left maxil- 
lula, in ventral aspect. (Scales for Figs. 29-32 all = 0.5 mm.), b. Sternal surface. 
ex. Exopodite. /. Paragnath. p. Mandibular palp. u. Upper lip or labrum. 4. Segment 
of endopodite distal to " knee ". 



ON A STYGIOMYSJS FROM THE WEST INDIES 315 

pairs are modified as maxilliped and gnathopods ; the same is true of the Eucopiidae, 
but here the following three pairs (5-7) are long and prehensile (see pp. 305 and 319). 
This brings up the question of the terminology of the thoracic appendages, which 
is discussed in the next section. 



IV. TERMINOLOGY OF THE THORACIC APPENDAGES. 

(a) Terminology of the appendages. Various names have been given to the 
thoracic appendages of Malacostraca according to the special functions for which 
they may be adapted. Dr. O. S. Tattersall writes of the Mysidacea : " since different 
appendages may be adapted for this or that function in different genera, I consider 
such terms most confusing, and in this work I propose to use one term, thoracic 
limbs, for these eight appendages, and to refer to each under its appropriate number, 
irrespective of the particular function for which it has become adapted." (Tattersall 
& Tattersall, 1951, pp. 18-19.) That there is some confusion is apparent if one 
examines certain statements in the joint work just cited. For example, under the 
definition of the order Mysidacea (pp. 11-12) " first, and sometimes the second, 
pairs modified as gnathopods or maxillipeds " might be interpreted in different 
ways — that both pairs are either gnathopods or maxillipeds, or that one pair is to 
be regarded as maxillipeds the other as gnathopods. On p. 24 " second thoracic 
limb (= gnathopod or 2nd maxilliped) " is also ambiguous — are the terms used 
synonymously or not ? On p. 97 " endopods of the second to the fourth pairs of 
thoracic limbs . . . developed as gnathopods " implies that the first pair are 
maxillipeds. Most of the confusion centres round these two terms. Sometimes 
they are used synonymously ; sometimes each is used in a narrower sense, the 
anterior pair (or pairs) which are more markedly modified for " Nahrungszerkleine- 
rung " being called maxillipeds in contrast to the less markedly modified succeeding 
pair (or pairs) of gnathopods (see Zimmer, 1927, pp. 302-303). 

Nor is there agreement as to the number of pairs of thoracic appendages which 
are modified as maxillipeds in the various orders of Malacostraca (the term gnathopod 
is commonly used in literature dealing with the superorder Peracardia, rarely if 
at all in that dealing with Eucarida, Hoplocarida or Syncarida). In primitive 
types none of the thoracic limbs are differentiated as maxillipeds (Caiman, 1909, 
p. 145) ; this is the case in Nebaliacea and Euphausiacea according to most authors. 
In other orders one, two, or three pairs are generally regarded as maxillipeds. With 
regard to the Stomatopoda Caiman (1909, p. 322) wrote : " The first five pairs of 
thoracic appendages are similar in structure and are commonly called maxillipeds, 
though, as they possess no endites or other adaptations for mastication, the name 
is hardly appropriate ". Yet it is still often said that the Stomatopods have five 
pairs of maxillipeds — here the term gnathopod might be more appropriate. Hansen 
(1925, pp. 94, 128 and 154) thought that throughout the whole of the Malacostraca 
there is one pair of real maxillipeds. In his summing up (p. 154) he says : " Maxil- 
lipeds and thoracic legs. — . . . They are similar in Leptostraca, while in the other 
orders the first pair, the real maxillipeds, differ somewhat or considerably or very 
much from the following pairs, the legs." Yet even he inadvertently introduces 



3 i6 ON A STYGIOMYSIS FROM THE WEST INDIES 

further confusion because : (i) the reader has to bear in mind that his thoracic legs 
1-7 are in reality thoracic appendages 2-8 and (ii) under the Decapoda he has 
reverted to the usual terminology of maxillipeds 1-3 and peraeopods or walking 
legs 1-5 (= thoracic limbs 4-8). Also under the Mysidacea he speaks of " inter- 
mediate " legs. 

The terminology of the thoracic limbs has recently been discussed by Siewing 
(1958, pp. 250-251). The maxillipeds, he says, are chiefly adapted for chewing or 
filtering food and are characterized above all by the presence of endites. " Sind die 
Endite deutlich grosser als die den nach hinten folgenden Extremitaten, so wird 
man sie betreffenden Extremitaten als Maxillipeden von den folgenden differenzieren. 
In diesem Sinne ist das 1 Thorakalbeinpaar, zum Teil auch das zweite, der Mysidacea 
als Maxilliped zu bezeichnen, denn es besitzt an der Basis einen abgegliederten 
Endite ; Kaufortsatze werden ausserdem an Praeischium und Ischium beobachtet." 
Gnathopods, on the other hand, have he says their distal segments modified, often 
to form chelae or subchelae ; but according to Tattersall & Tattersall (195 1, p. 24) 
the first thoracic limbs of Mysidacea may sometimes be subchelate. 

Siewing's statement concerning the Mysidacea agrees in part with that of Caiman 
(1909, p. 174) : " The first pair of thoracic limbs are always specialized as maxillipeds. 
In the Lophogastridae and Eucopiidae they are without distinct endites (italics mine). 
In the Mysidae an endite is generally borne on the basipodite, and sometimes also 
by each of the two following segments." Tattersall & Tattersall (1951, p. 97) also 
say that in Eucopia there are " no gnathobasic lobes on second, third or fourth 
segments ". If Siewing's definitions are strictly applied, the first thoracic limb of 
Eucopia, though smaller than, and appreciably different from, each of the three 
succeeding pairs of appendages, would also have to be called a gnathopod (see 
Text-fig. 31 of this paper) . The same applies to the first thoracic limb of, e.g. Mysidopsis 
which also has no endites (gnathobasic lobes) on second, third or fourth segments 
(Tattersall & Tattersall 1951, p. 316) and to that of Stygiomysis although it is much 
smaller, and more intimately associated with the mouth parts, than are the three 
following pairs (Text-figs. 7, 9 and 10). Caroli (1937, pp. 22 and 226), who refers 
to the first pair as maxillipeds, gives the neatest possible definition of the next three 
pairs with his " Toracopodi 2-4 (gnatopodi) massillipediforme "! Typhlolepidomysis, 
on the other hand, exemplifies the two categories exceptionally well, having large 
endites on the first pair which are not pediform, none on the pediform second pair 
which have strong spines on the more distal segments. Unfortunately, Villalobos 
(1951, p. 205) refers to the latter as " primer par de pereiopodos " in the text and 
as " maxilipedo del secundo par " in the legend to Text-fig. 10. In Lepidops the 
second pair differs but slightly from the following pairs of thoracic limbs. In the 
family Petalophthalmidae the second thoracic limb has a large lamellar expansion 
on the ischium (merus of some writers) which, with its partner, forms a floor to the 
whole mouth region. (I have seen a specimen which had been preserved while 
enjoying a meal, and these lobes apparently help to retain the food mass while it is 
being chewed). Should this appendage be termed a maxilliped? It is much larger 
than the first thoracic limb (which in Hansenomysis has no endites) and its distal 
segments are modified to assist in manipulating the food mass. It seems illogical 



ON A STYGIOMYSIS FROM THE WEST INDIES 



3i7 



to term the first thoracic appendage a maxilliped in Mysis but not in Mysidopsis, 
in Petalophthalmus but not in Hansenomysis, in Gnathophausia (Text-figs. 33 and 
34) but not in Lophogaster — see p. 316. Form depends on function and the feeding, 
locomotor and other habits of the animals determine how many appendages will 
be modified, and to what extent. The requirements of a raptorial animal differ 




Figs. 33-34. Gnathophausia ingens (Dohrn). (Unregistered specimen from Madeira). 
Fig. 33. Right maxilliped, in ventral aspect. Fig. 34. Part of same, in dorsal aspect. 
e. Endite. ep. Epipodite. ex. Exopodite. 3 and 4. Segments of endopodite on either 
side of " knee " joint. 



3* 



ON A STYGIOMYSIS FROM THE WEST INDIES 



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ON A STYGIOMYSIS FROM THE WEST INDIES 319 

from those of a filter feeder, even if the latter can also manipulate large food masses. 
The habitat to which an animal is adapted, even the niche within the habitat, may 
lead to modifications in both filtratory and raptatory feeders. Eucopia, with its 
long, slender " prehensile " endopodites 5-7, may remain suspended in mid water 
and seize any food that happens to come within range of these long legs, passing 
it on to the short gnathopods (2-4) ; it may even pursue its prey since the pleopods 
are well developed and natatory (see also p. 305). I fully agree with Siewing that 
terms ought to be properly defined, and that the terms under discussion need re- 
defining. But, for the reasons given above, I do not think that one can apply his 
definitions too rigidly, within the order Mysidacea at any rate. 

Cannon (1927) has shown that all the thoracic limbs in Nebalia are specially 
adapted for a unique mode of filter feeding. More recently Barkley (1940) has 
figured in great detail thoracic limbs 1-6 of Euphausia and has found that these 
six pairs together form an elaborate basket or filter for straining minute organisms 
from the water (the last two pairs of thoracic appendages are greatly reduced). 
In both these genera the similarity of the thoracic limbs would appear to be a second- 
ary specialization rather than a primitive character. 

(b) Terminology of the segments of the endopodite : There is also consider- 
able diversity of opinion regarding the terminology of the segments in the endopodite 
of the thoracic appendages. In Stygiomysis holthuisi, for example, it is possible 
to interpret the parts numbered 1-6 in Text-figs. 9, 10 and 11 in three different 
ways : (i) ischium, merus, carpus, propodus, dactylus, and claw ; (ii) preischium, 
ischium, merus, carpus, propodus, and dactylus (with which the claw is incorporated); 
(iii) preischium, ischium, merus, carpo-propodus, dactylus, and claw. According 
to (i) the " knee " joint is at the junction of carpus and propodus ; according to 
(ii) and (iii) it lies between merus and carpus. This is why I have simply numbered 
the segments of the endopodite — those distal to the basis which bears the exopodite 
and is part of the protopodite. 

The different terminologies which are in use are best shown in tabular form (Table 
I, which may not be complete). To the student reared on Caiman's text book 
terminology A will be familiar ; according to this the " knee " joint is sometimes 
between merus and carpus, sometimes between carpus and propodus. The other 
terminologies (B, C and D) we owe to Hansen (1925). He maintains that there is 
an additional segment, the preischium, which is distinct in Peracarida but is often 
fused with the ischium (Eucarida) or with the basis (Syncarida) . He also maintains 
that throughout the Eumalacostraca the " knee " is at the junction of merus and 
carpus and that, where there appears to be only two segments distal to the " knee ", 
the carpus is fused with the propodus (the claw is not a true segment and is not 
included). As shown in the table, this fusion of carpus and propodus occurs very 
frequently in Mysidacea and throughout the other Peracaridan orders. Siewing 
(1958) has recently studied Thermosbaena and his interpretation of the segmentation 
is shown as Ei and E2. In minute animals like Thermosbaena and the Bathynellacea 
there is a reduction in the number of segments and it is by no means easy to decide 
where the " knee " is. Siewing thinks that here there is no preischium, and that the 
ischium is either fused with the basis or more or less completely fused with the merus. 



3 20 ON A STYGIOMYSIS FROM THE WEST INDIES 

In Stygiomysis holthuisi the number of segments in the endopodite is identical 
in each of the eight pairs of thoracic limbs (Text-figs. 7, 9-11). In each there are 
three segments proximal to the " knee " and three distal to it (including the claw 
or claw-like dactylus, according to the terminology used). Hansen would doubtless 
have regarded segment 4 as a fused carpo-propodus (B4) rather than as the carpus 
followed by the propodus and claw-like dactylus (Bi). In Tattersall & Tattersall 
(1951, p. 24) it is stated that, in both first and second pairs of thoracic limbs of 
Mysidacea, carpus and propodus are always fused to form a single segment. Yet 
in fig. 42, C on p. 199 the first thoracic endopodite of Erythrops elegans (G. O. Sars) 
is represented as having four segments proximal to the " knee " joint and two plus 
claw distal to it. This can only be interpreted (if correct and not a slight lapse on 
the part of the artist) as : preischium, ischium,, merus, carpus, [" knee "], propodus, 
dactylus and claw. According to Hansen the " knee " joint in Peracarida does not 
come at the junction of carpus and propodus. Since fig. 42, D shows the second 
endopodite of Erythrops elegans with only three segments proximal to the " knee ", 
fig. 42, C may not be correct. Hansen (1925) based his interpretation of the segmenta- 
tion, in part at least, on a study of the muscles (see his figures of the distal segments 
of the thoracic endopodite in Mysidacea on pi. vi). In Stygiomysis there appear to 
be no muscle bands in segment 5, which might support Hansen's interpretation 
of 5 and 6 being dactylus and claw respectively. Perhaps the time has come for 
a thorough reappraisal of Hansen's work along the lines of the detailed functional 
analysis of limb structure undertaken in recent years by Dr. Manton in terrestrial 
Arthropoda (Manton, 1950-58 — a series of papers on the Evolution of Arthropodan 
Locomotory Mechanisms ; the work is still in progress). 



V. ADDITIONAL NOTE ON SPELAEOGRIPHUS LEPIDOPS GORDON 

Order SPELAEOGRIPHACEA Gordon, 1957 

Spelaeogriphacea Gordon, 1957, P- 45- 

Spelaeogriphacea, Siewing, 1958, p. 267 ; Kaestner, 1959, p. 918. 

" Reihe " Spelaeogriphacea Siewing, 1959, p. no, Fig. 65. 

Systematic Position. Siewing (1958, p. 247) placed the order Spelaeogriphacea 
among those Eumalacostraca with the telson and last abdominal somite fused. 
But, as stated in my original description and as shown in PL 4 of this paper, the 
sixth pleonite is free from, and much larger than, the telson. In a later work, which 
has just appeared, Siewing (1959, p. no, fig. 65 — cited in his 1958 paper as if it 
had already been published) has raised the Spelaeogriphacea to the rank of a "Reihe" 
or superorder, without apparently stating his reasons for doing so. 

Since Spelaeogriphus, the only known genus, undoubtedly possesses a Peracaridan 
brood pouch composed of oostegites, I still regard the Spelaeogriphacea as an order 
belonging to the superorder (" Division " of Caiman) Peracarida. Its precise relation- 
ship to the other Peracaridan orders cannot be determined until the internal anatomy 
and the embryology of Spelaeogriphus are known. However, from its external 



ON A STYGIOMYSIS FROM THE WEST INDIES 321 

morphology, it appears to have affinities with the Tanaidacea, since it has a small 
but distinct carapace and a pair of cup-shaped respiratory organs beneath the cara- 
pace, near the bases of the first thoracic limbs (the maxillipeds). Siewing (1956, 
p. 8y) has pointed out that in the Tanaidacea " der hinter dem Maxillipeden an 
der Korperwand liegende loffelartige Fortsatz ist kein Epipodit, wie Dohrn bei 
seinen embryologischen Untersuchungen zeigen konnte." (Dohrn, 1870, pi. xi). 
The respiratory organ in Spelaeogriphus may also be of this kind and not a true 
epipodite, but I called it an epipodite since that is the term commonly used in 
systematic descriptions of Tanaidacea. 

However, I could not include Spelaeogriphus in the order Tanaidacea, since it 
differs in many respects from that group, having certain affinities with the Syncarida 
and with Monodella and some which are unique. In his earlier paper Siewing (1957, 
p. 267, phylogenetic diagram) placed it within the Peracarida, provisionally near 
the Isopoda-Tanaidacea line. Unless Dr. Siewing has studied material of Spelaeo- 
griphus lepidops and has found that it differs markedly, anatomically and /or 
embryologicall y , from the other Peracaridan orders, his decision to separate Spelaeo- 
griphacea from the " Reihe " Peracarida is premature, and in my opinion this step 
is unnecessary. It is a pity that this alteration was made in a text book because in 
all probability his phylogenetic scheme will be reproduced in future text books. 
The one thing that all Peracaridan orders have in common is that the young are 
incubated until a late stage of development in a ventral thoracic brood pouch or 
marsupium ; Siewing's (1958) separation of the order Thermosbaenacea from the 
true Peracarida is much more reasonable, since in both the known genera there is a 
dorsal carapacial marsupium. Stella (1959, p. 122) has accepted the " Reihe " 
Pancarida although she unfortunately refers to it as a " phylum ". Raising the Spelae- 
ogriphacea to superordinal rank does not solve the problem of its relationships. 
The greatest caution should be exercised in establishing higher categories like 
superorders and subclasses until the animals have been thoroughly studied. 



Genus SPELAEOGRIPHUS Gordon, 1957 

Spelaeogriphus Gordon, 1957, P- 3 2 - 
Spelaeogriphus, Kaestner, 1959, p. 918. 



Spelaeogriphus lepidops Gordon, 1957 

Spelaeogriphus lepidops Gordon, 1957, p. 33, Text-figs. 1-26. 
Spelaeogriphus lepidops, Kaestner, 1959, p. 918, Text-fig. 816. 

Additional material. Bats Cave, Table Mountain, 30.ix.56, six specimens 
fixed in Bouin's solution (4 <J, 1 ovigerous and 1 smaller $). From a stream at bottom 
of Bats Cave (temp, of water 50-5° F.), i.ix.57, seven specimens (5 <$, 1 ovigerous 
? and 2 immature specimens). 

Remarks. The specimens were collected by the South African Spelaeological 



322 ON A STYGIOMYSIS FROM THE WEST INDIES 

Association ; the 1957 material was fixed in 96 per cent ethyl alcohol (97 parts) and 
glacial acetic acid (3 parts). This fixative seems to be too drastic, causing the speci- 
mens to curl up to such an extent that in one instance the uropods got entangled 
among the thoracic appendages bunched up beneath the carapace region. 

I have had several requests for a good figure of the entire animal, suitable for 
reproduction in a text book. My original Text-fig. 1 was incomplete, since I did not 
include the appendages, and Kaestner (1959, fig. 816) has based his figure on Dr. 
Barnard's sketch which I reproduced as Text -fig. 26 of my 1957 paper. The anten- 
nulae and antennae are not represented and these alter the look of an animal appre- 
ciably ; in Kaestner's reproduction the first three thoracic exopodites look not unlike 
the gills of Lophogastrid Mysids. I take this opportunity, therefore, of publishing an 
original drawing of the whole animal, an ovigerous female, in dorso-lateral aspect 
as PI. 4. This figure will also illustrate my article on Spelaeogriphacea which 
will appear in due course in the McGraw-Hill Encyclopedia of Science and Tech- 
nology. 

The female represented in PL 4 measures 6 mm. in length, from rostral apex 
to tip of telson, and the antenna is almost or quite as long as the body. The two 
antennular flagella are slightly unequal (they are more so in the male) ; the inner 
flagellum has about 30, the outer one about 24, segments. The faint line down the 
right side of the body, from carapace to first abdominal somite, is an artefact, a 
slight fold or crease in the integument. The marsupium contains ten to twelve 
rather large ova. I have not attempted to straighten out the thoracic limbs, which 
in preserved material tend to bend inwards beneath the body, because they easily 
break off if touched. 

There is not much to add to my original description, since the animals do not 
seem to grow much larger ; the largest male measures 8-6 mm. in length. Its antenna 
is at least 8 mm. long ; the inner flagellum of the antennula is appreciably longer than 
the outer one and the number of segments in each is about 45 and 26 respectively. 

In my original description I used Caiman's terminology for the various segments 
of the thoracic limbs (see Table I, A). But, according to Hansen (1925), there is 
always a distinct preischium in the Peracarida (Table I, Bi to B4). The segments 
of the endopodite would thus be termed : preischium, ischium, merus, [" knee " 
joint], carpo-propodus and dactylus (Table I, B4). I have re-examined these 
appendages and in some there seems to be a faint indication of separation of the 
claw from the dactylus. In one specimen the distal sharp point is broken off neatly, 
beyond the longest spine on the ventral margin ; this part doubtless represents 
the claw or nail (Gordon, 1957, Text-figs. 19 and 21-25). 

The entire length of the gut is filled with a dark brown or blackish mass, suggesting 
that the animal feeds largely on detritus. 

The registration numbers of the type specimens in the British Museum Collection 
are : holotype ((J), 1957.xi.12-i and allotype (?), 1957. xi. 12.2. 

I have a correction to make : in my 1957 paper (p. 43) I said that the Tanaidacea 
are entirely marine. But I overlooked a paper by Stephensen (1936) in which he 
described Tanais stanfordi from a freshwater lake in the Kurile Islands and mentions 
at least one other freshwater species from South America. 



ON A STYGIOMYSIS FROM THE WEST INDIES 323 

LIST OF WORKS REFERRED TO. 

Allaud, Ch. & Jeannel, R. 1914. In Jeannel, R. and Racovitza, E. G. Enumeration des 

grottes visitees 1911-1913. (5) 413. Arch. Zool. exp. gen. (Biospeologica xxxiii) 53 (7) : 

325-558> 5° text-figs. [p. 381]. 
Barkley, E. 1940. Nahrung und Filterapparat des Walkrebschen Euphausia superba Dana. 

Z.f. Fisch. Beiheft 1, Walforschung 1, Enhalt IV : 65-156, 39 text-figs. 
Bolivar Y Pieltain, C. 1943-44. Exploracion biologica de algunas cavernas de Cuba. Ciencia 

Mexico. 4 (11-12) : 301-304. 

1950. Estudo de una Cirolana cavernicola nueva de la region de Vallas, san Luis Potosi, 

Mexico. Ciencia Mexico 10 (7-8) : 21 1-2 18. 

Calman, W. T. 1909. A Treatise on Zoology, edited by Sir Ray Lankester, Part 7, Appendi- 

culata, Fasc. 3 Crustacea. London (A. & C. Black) : viii + 346 pp. Text-figs. 
1932. A cave-dwelling Crustacean of the family Mysidae from the Island of Lanzarote. 

Ann. Mag. nat. Hist. (10) 10 : 127-131, 6 text-figs. 
Cannon, H. G. 1927. " On the feeding mechanism of Nebalia bipes." Trans. R. Soc. Edinburgh. 

55 : 355-37°, 7 text-figs. 
Cannon, H. G. & Manton, S. M. 1927. On the feeding mechanism of a Mysid Crustacean, 

Hemimysis lamomae. Trans. R. Soc. Edinburgh, 55 : 219-253, 4 pis., 16 text-figs. 
Caroli, E. 1924. Su di un misidaceo cavernicolo (Spelaeomysis bottazzii n.g., n.sp.) di terra 

d'Otranto. Atti Ace. Naz. Lincei Roma, 33 (2) : 512-513. 

1937- Stygiomysis hydruntina n.g., n.sp., Misidaceo cavernicolo di Terra d'Otranto, 

rappresentante di una nuova famiglia. Nota preliminare. Boll. Zool. Torino, 8 : 219-227, 
3 text-figs. 

Chappuis, P. A. i 93 i. Copepoda Harpacticoidea der Deutschen limnologischen Sunda- 

Expedition. Arch. Hydrobiol. Stuttgart. Suppl. 8 : 512-584, 152 text-figs. 
Creaser, E. P. 1936. Crustaceans from Yucatan. Publ. Carnegie Inst. Washington, 457 : 

1 17-132. 43 text-figs. 
— — 1938. Larger cave Crustacea of the Yucatan Peninsula. Publ. Carnegie Inst. Washington, 

491 : 159-164, 8 text-figs. 
Dohrn, A. 1870. Untersuchungen fiber Bau und Entwicklung der Arthropoden. VII. Zur 

Kenntnis vom Bau und der Entwichlung von Tanais. Jenaische Zeitschr. Nat. 5 : 293-306, 

pis. 11 and 12. 
Fage, L. 1924. Sur un type nouveau de Mysidace des eaux souterraines de Tile de Zanzibar. 

C.R. Acad. sci. Paris. 178 : 2127. 

1925. Lepidophthalmus servatus Fage. Type nouveau de Mysidace des eaux souterraines 

de Zanzibar. (Biospeologica 51). Arch. zool. Exper. Gen. 63 : 525-532, 7 text-figs. 

1941. Mysidacea : Lophogastridae — I. Dana Rept. Copenhagen, 19 : 1-52, 51 text-figs. 

1942. Mysidacea : Lophogastridae — II. Dana Rept. Copenhagen, 23 : 1-67, 42 text-figs. 

Gordon, I. 1957. On Spelaeogriphus, a new cavernicolous Crustacean from South Africa. 

Bull. Brit. Mus. (Nat. Hist.), Zool. 5 (2) : 31-47, 26 Text-figs. 
1958. A new subterranean Crustacean from the West Indies. Nature, Lond. 181 : 1552-1553, 

2 text-figs. 
Hansen, H. J. 1925. Studies on Arthropoda. Copenhagen. Vol. II, 176 pp., 8 pis. 
Kaestner, A. 1959. Lehrbuch der Speziellen Zoologie. Pt. I : Wirbellose, Lief. 4. Jena. 

pp. 659-979, text-figs. 630-861. 
Manton, S. M. 1928. On some points in the anatomy and habits of the Lophogastrid Crustacea. 

Trans. R. Soc. Edinburgh, 56 : 1 03-1 19, 3 pis., 2 text-figs. 
1950-1958. The Evolution of Arthropodan Locomotory Mechanisms. Parts 1-6. /. Linn. 

Soc. (Zool.) London, 41 : 529-570, 4 pis. 8 text-figs.; 42 : 93-117, 5 text-figs.; 42 : 1 18-166, 

12 text-figs., 6 pis.; 42 : 299-368, 4 pis., 8 text-figs.; 43 : 153-187, 1 pi., 8 text-figs.; 

43 : 487-556, 1 pi., 31 text-figs. 
Monod, Th. 1940. Thermosbaenacea in Bronn's Klassen und Ordnungen des Tierreichs, 

5, 1 (4) : 1-24, 32 text-figs. Leipzig. 



3 2 4 ON A STYGIOMYSIS FROM THE WEST INDIES 

Nicholls, G. E. 193 1 . Micraspides calmani, a new Syncaridan from the West Coast of Tasmania. 

/. Linn. Soc. Lond. 37 : 473-488, 2 pis., 1 text-fig. 
Nouvel, H. 1943. Mysidaces provenant des campagnes du Prince Albert ier de Monaco. 

Res. Camp. sci. Monaco, 105 : 1-128, 180 text-figs., 5 pis. 
Sars, G. O. 1885. Report on the Schizopoda collected by H.M.S. Challenger during the years 

1873-76. Zool. Challenger Exp. 13, Pt. 37 : 228 pp., 38 pis., text-figs. 
Sars, M. 1862. Beskrivelse over Lophogaster typicus, en maerkvaerdig form af de lavere 

tifoddede Krebsdyr . . . Universitets-program . . . pp. iv and 37, 3 pis. 4 Christiania 

[With resume in French]. 
Siewing, R. 1958. Anatomie und Histologic von Thermosbaena mirabilis. Ein Beitrag zur 

Phylogenie der Reihe Pancarida (Thermosbaenacea). Akad. Wiss. und Liter atur. Abh. 

Math. -Nat. Kl. Jahr. 1957 No. 7 : 197-270, 43 text-figs. 
1959- Syncarida in Bronns Klassen und Ordnungen des Tierreichs. V, 1, 4 (2) : 1-121, 

65 text-figs. 
Stammer, H. J. 1933. Einige seltener oder neue Hohlentiere. Verh. Deutsch. zool. Ges. Leipzig. 

35 : 263-266. 
1936. Ein neuer Hohlenschizopode, Troglomysis vjetrenicensis n.g., n.sp. . . . Zool. Jahrb. 

Jena (Syst.) 68: 53-104, 30 text-figs. 
Stella, E. 1959. Ulteriori osservazioni sulla riproduzione e lo sviluppo di Monodella argentarii 

(Pancarida, Thermosbaenacea). Rivista di Biologia 51 (1) [n.s. vol. xi] : 1 21-144, ° text- 
figs., 2 pis., 1 table. 
Stephensen, K. 1936. A Tanaid (Tanais stanfordi Richardson) found in fresh water in the 

Kurile Islands . . . Annot. Zool. Japon. 15 (3) : 361-373, 5 Text-figs. 
Tattersall, O. S. 1957. Report on a small collection of Mysidacea from the Sierra Leone 

estuary, together with a survey of the genus Rhopalophthalmus Illig and a description of 

a new species of Tenagomysis from Lagos, Nigeria. Proc. zool. Soc. Lond. 129 : 81-128, 15 

text-figs. 
Tattersall, W. M. 1939. The Euphausiacea and Mysidacea of the John Murray Expedition 

to the Indian Ocean. Sci. Rep. John Murray Exped. London, 5 : 203-246, 21 text-figs. 
1951. A review of the Mysidacea of the United States National Museum. Bull. U.S. nat. 

Mus. 201 : 292 pp., 103 text-figs. 
Tattersall, W. M. & Tattersall, O. S. 1951. The British Mysidacea. Ray Society London 136 : 

pp. viii + 460, frontispiece and 118 text-figs. 
Villalobos, A. 1951. Un nuevo misidaceo delas Grutas de Quintero en el Estado de Tamaulipas. 

An. Inst. Biol. Mexico, 22 : 191-128, 14 text-figs. 
Zimmer, C. 1927. Allgemeine Einleitung in die Naturgeschichte der Crustacea (pp. 277-304, 

text-figs. 216-242) ; Mysidacea (pp. 607-650, text-figs. 592-645) in : Kukenthal und 

Krumbach, " Handbuch der Zoologie " 3 (1). Berlin (de Gruyter) 4 . 



PLATE 3 

Stygiomysis holthuisi (Gordon). Holotype, in dorsal aspect, a. Biramous antennula. b. 
Antenna, o. Ocular scale. 5. Fifth thoracic somite. 1 and 6. First and sixth abdominal somites. 

PLATE 4 
Spelaeogriphus lepidops Gordon. Ovigerous female in dorso-lateral aspect, a. Biramous an- 
tennula. b. Antennal flagellum. g. Respiratory exopodites or gills. 0. Ocular scale, s. Antenna! 
scale or exopodite. 5. Fifth thoracic somite. 1 and 6. First and sixth abdominal somites. 



♦*> 



Bull. B.M. (N.H.) Zool. 6, 5 



PLATE 3 




Bull. B.M. {N.H.) Zool. 6, 5 



PLATE 4 




CONTRIBUTIONS TOWARD A 

CLASSIFICATION OF MODERN 

ISOSPONDYLOUS FISHES 




WILLIAM A. GOSLINE 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ZOOLOGY Vol. 6 No. 6 

LONDON: i960 



CONTRIBUTIONS TOWARD A CLASSIFICATION 
OF MODERN ISOSPONDYLOUS FISHES 



BY 

WILLIAM A. GOSLINE 




Pp. 325-365 ; 15 Text-figures 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY Vol. 6 No. 6 

LONDON : i960 



THE BULLETIN OF THE BRITISH MUSEUM 
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issued in five series corresponding to the Departments 
of the Museum, and an Historical Series. 

Parts will appear at irregular intervals as they become 
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within one calendar year. 

This paper is Vol. 6, No. 6 of the Zoological series. 



© Trustees of the British Museum, i960 



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Issued April, i960 Price Fifteen Shillings 



CONTRIBUTIONS TOWARD A CLASSIFICATION 
OF MODERN ISOSPONDYLOUS FISHES 

By WILLIAM A. GOSLINE 

That the classification of the modern isospondylous fishes (Order Clupeiformes) 
is unsatisfactory would be admitted by all who have worked with them. The present 
contributions to the subject are divided into four sections. The first traces one 
structural complex, the caudal skeleton, through the whole group. The second 
and third have to do with controversial sections of isospondylous classification. 
In the final section a revised arrangement of modern forms, down to superfamily, 
is presented and its rationale discussed. The sections are in logical sequence, but 
it may also be noted that they are arranged in order of descending assurance on 
the part of the author. More, perhaps, than usual with classifications, that presented 
in section four is merely a " progress report ". 

Regarding nomenclature, the ordinal, subordinal, and superfamily names are 
formed according to the system proposed by Berg (1940 : 353). The following 
procedures concerning common names have been adopted. Where an old literature 
name is well known it will be used. Thus the members of the order Clupeiformes 
will be called the isospondylous fishes, and those of the suborder Esocoidei the haplo- 
mous fishes. Where a common name in this paper ends in old, it refers to a member of 
a suborder. For example, the term salmonoid fishes will be used to designate all 
of the forms of the Suborder Salmonoidei and not just those of the superfamily 
Salmonoidae. No common names will be used for superfamily 1 . 

Bone nomenclature is that of Devillers (1958). 

The work on which this paper is based was nearly all done at the British Museum 
(Natural History) and I wish to thank the members of the fish division of that 
Museum for the facilities extended me and for permission to examine the skeleton 
and fish collection. I also wish to thank them, notably N. B. Marshall, for the discus- 
sion and suggestion of various points incorporated into the paper. The following 
individuals have been kind enough to read part or all of the manuscript and to 
offer constructive criticism : E. H. Ahlstrom, H. B. Bigelow, D. M. Cohen, D. H. 
Dunkle, N. B. Marshall, and G. W. Mead. Finally, I am greatly obligated to the 
John Simon Guggenheim Foundation for the fellowship that has made this work 
possible. 

1 If Berg's termination oidei is used for suborder, the problem of devising different common names for 
subordinal and superfamily categories is equally great whether the superfamily ending oidea, recom- 
mended by the London Colloquium, or Berg's oidae is used. 

ZOOTL. 6, 6. 2I§ 



328 A CLASSIFICATION OF MODERN ISOSPOND YLOUS FISHES 

SECTION i. THE CAUDAL SKELETON OF 
MODERN ISOSPONDYLOUS FISHES 

INTRODUCTION 

The caudal skeleton of teleostean fishes has been the subject of papers by Barring- 
ton (1937), Blanco (1935), Hollister (1936, etc.), Kolliker (i860), Lotz (1864), Regan 
(1910a and b), Totton (1914), Whitehouse (1910a and b, 1918), and others. Never- 
theless, the accumulated information is rather amorphous, and the caudal skeleton 
is usually either ignored or briefly dismissed in taxonomic papers. A large part of 
the difficulty would seem to stem from the failure to establish stable reference 
points as a basis for comparing one caudal skeleton with another : hence the intro- 
ductory remarks that follow. 

In a relatively simple caudal skeleton like that of Amia (Goodrich, 1909, fig. 311 
or Whitehouse, 1910a, fig. 5), each of the last several vertebrae articulates with the 
base of a single caudal ray by means of a hypural (which is probably a hemal spine 
fused with the radial element of the fin ray, cf., Totton, 1914 : 255). In the teleostean 
fishes, however, complications of several types occur. In the first place, several 
of the posteriormost vertebral centra disappear (see below). Second, the hypural 
elements themselves become reduced in number, presumably through loss and fusion, 
so that in teleosts there are usually two to several caudal rays articulating with 
each hypural. Third, one or more elements of the caudal skeleton above the vertebral 
column lose their proximal contact with the neural arches (this is also true of Amia) 
becoming the so-called epurals. Finally, in the more primitive isospondylous fishes 
there are one to four pairs of bones that lie along the posterior end of the spinal 
cord ; these are the uroneurals. 

A final term, urostyle, has been used in the literature to cover such a wide variety 
of structures that it will be completely avoided here. Supposedly the urostyle 
represents the last upturned vertebral centra. However, Regan (1910a) and Ramanu- 
jam (1929) have shown that the so-called urostyle of the Clupeidae is mostly formed 
of uroneurals (see footnote 2 of the present paper), and the uroneurals may form 
a large part of the " urostyle " of at least some higher teleosts. 

Another difficulty lies in the fact that at least two of the centra at the end of the 
vertebral column in such a fish as Elops disappear as separate entities in the higher 
teleosts. Thus the last visible centrum of one fish is not always the homologue of 
the last centrum of another. To get around this difficulty it is necessary to establish 
some reference point that is both homologous and identifiable in as many teleosts as 
possible. The most satisfactory seems to be the posteriormost vertebra that is 
consistently present in all isospondylous fishes (except for such aberrant forms as 
Coilia) and is usually present in other teleosts ; this is here termed the terminal 
vertebra (TV of Text-figs. 1-15). It can usually be identified by a series of criteria 
(though the misimpression should not be given that any or all of these will easily 
or even securely establish the terminal vertebra of all fishes). The best single feature 
is probably the nature of the anteriormost hypural articulating or fused with it. 

In most lower teleosts (and indeed in many percoids) the hypurals to the lower 
caudal lobe are quite constant in number (three) and general shape. The uppermost 



A CLASSIFICATION OF MODERN ISOSPON D YLOUS FISHES 329 

of these (HY3 of Text-figs.) is a rather narrow strut that is never expanded posteriorly. 
The next hypural below (HY2 of Text-figs.) is broadly wedge-shaped, i.e., consider- 
ably expanded toward the rear. Below this again is another strut-like hypural with 
a broad basal articulation or area of fusion with a centrum ; throughout the paper 
this will be considered the lowermost hypural (HYi of Text-figs.), and the vertebra 
with which it articulates is the terminal vertebra. This hypural, or hypural 1, 
almost always differs from those behind and above it in having on each side near its 
base a lateral flange, which usually terminates posterolaterally in a sharp point. (This 
flange serves for the attachment of part of the flexor caudalis ventralis superficialis 
muscle (Greene and Greene, 1914 : 46) and probably in part as protection for the 
blood vessels that exit from the vertebral column behind the base of hypural 1. 
In these respects hypural 1 is the terminal structure retaining a number of hemal 
arch characteristics.) Generally, the terminal vertebra may also be identified in 
other ways. Whatever its relationship with centra (if present) behind it, there is 
always a normal, hourglass-shaped centrum and a typical intervertebral articulation 
immediately in front of it. More diagnostic perhaps is that if the front pair of uron- 
eurals wedge into or fuse with any centrum, it is that of the terminal vertebra (except 
apparently in osteoglossoids) . 

In some recent teleosts one or two separately ossified centra may remain visible 
in the adult ; if so these are here termed postterminal 1 centra (PT of Text-figs. 1-4, 
6, 7, 10 and 13-15) and are numbered from front to rear. When there are two post- 
terminal centra, hypural 1 alone articulates or fuses with the terminal vertebra, 
and hypurals 2 and 3 arise together on the first postterminal centrum (as in Text-figs. 

1-4). 

Vertebrae just anterior to the terminal vertebra are here called preterminal 
vertebrae (PR of Text-figs.) and are numbered from back to front. 

The structure of the upper half of the caudal skeleton seems to be more subject 
to variation than the lower. This is perhaps to be expected as there is phylogenetic- 
ally a progressive retraction of the vertebral column from the upper caudal lobe. 
At any rate, in the isospondylous fishes there is some fluctuation, from specimen to 
specimen as well as from species to species, in the number of uroneurals and epurals, 
and from genus to genus in the number of upper hypurals. 

Two other types of ontogenetic and individual variation should be mentioned. 
One is the abnormal but frequent doubling of the neural or hemal arches attached 
to any one centrum, even the terminal centrum (Totton, 1914 : 253). The other is the 
nature of the association between these arches and their centra. In young fishes the 
neural and hemal arches are wedged into their centra and only later in ontogeny do 
they fuse with them ;• however, in some forms, e.g., Albula, such fusion never does 
take place. 

Finally, it should be noted that dried skeletons, which have been used here, 
show especially the superficial features of the bones whereas stained and cleared 
specimens show more of the deeper features. Furthermore, for practical reasons 

1 Dr. E. H. Ahlstrom informs me that he prefers to retain the term urostyle for those centra with 
which the hypurals articulate. If the confusion that has arisen from including the uroneurals in the 
" urostyle " can be cleared away, there is much to be said for this, especially in view of the term " post- 
terminal centra " with its self-contradictory adjective that is used in this paper. 



33° 



A CLASSIFICATION OF MODERN ISOSPON D YLOUS FISHES 



dried skeletons are usually prepared from large adults whereas stained and cleared 
specimens are preferably small. Thus an investigation based on dried material 
is likely to emphasize one set of caudal features, that based on cleared specimens 
another. 

COMPARATIVE CAUDAL MORPHOLOGY 

In the following accounts of individual caudal skeletons, a basal group (Diagram 
i) is dealt with first. The " lineages " of caudal structure presumably arising from 
these basal forms are then discussed. Finally the component members of these 



EPI EP2 



UN3 




Fig. i. Caudal skeleton of A lepocephalus rostratus, x 5 J. Stippling indicates interosseous 
or intraosseous areas. The arrow indicates the midpoint between the lobes of the caudal 
fin. EP, epural (these are numbered from top to bottom) ; HS, hemal arch and spine ; 
HY, hypural (hypurals are numbered from bottom to top ; hypural i is really in large 
part the hemal arch and spine of the terminal vertebra) ; NA, neural arch (only) ; NS, 
neural arch and spine ; PR, preterminal vertebra (these are numbered from rear to 
front) ; PT, postterminal centrum (theseare numbered from front to rear) ; TV, termi- 
nal vertebra ; and UN, uroneural (these are numbered from front to rear). 



" lineages " are taken up. Superfamily and subordinal names are those adopted in 
Section 4 of this paper ; for the moment it need only be noted that Chanos, Kneria, 
Phractolaemus and Cromeria are here included in the Gonorhynchoidei. Failure to 
mention certain families is due to lack of available material. 

Of the caudal skeletons examined the following would seem to be among the most 
primitive 1 : Elops (Hollister, 1936, fig. 14), Alepocephalus (Text-fig. 1), Salvelinus 
(Text-fig. 2), Esox (Text-fig. 3), and Hiodon (Text-fig. 4). All of these have a number 

1 Underlying the use of the terms " unspecialized " and " primitive " throughout this paper is the 
assumption that in teleostean fishes there has been a progressive fusion and loss of parts in the evolution 
of the caudal skeleton. (According to this concept the most highly simplified caudal skeletons are often 
the most " advanced " whereas the more complex are frequently the most " primitive ".) 



A CLASSIFICATION OF MODERN ISOSPOND YLOUS FISHES 



33i 



of unspecialized 1 features in common : the front end of the anterior uroneural 
overlaps, but neither wedges deeply into nor fuses with, the terminal vertebra ; 
hypural 1 is the only hypural articulating or fusing with the terminal vertebra ; 




Fig. 2. 



Caudal skeleton of Salvelinus grayi 

are included to show gap for posterior uroneurals 



X 6§. Bases of some of the upper fin rays 



hypurals 2 and 3, and these only, articulate with the first postterminal centrum ; 
and a portion of a second postterminal centrum is always visible. 

From the type of caudal skeleton represented in these primitive forms the more 
specialized caudal structures found among isospondylous fishes have presumably 
arisen. Indeed it seems possible to trace in the caudal skeleton four main " lineages " 
(I-IV of Diagram 1), as will be done here. As to these, Elops and Alepocephalus 



One or no postterminal 
centra in the adult 

Anterior uroneural either 
fused with or wedged 
into terminal vertebra 




Mormyriformes 
(Fig 15) 



Osteoglossoidae 
(Figs. 13, 14) 



Hypurals to the upper half of the 
caudal fin fused into a single 
plate-like structure 



Two postterminal centra, 
hypurals 2 and 3 articu- 
lating with postterminal 
centrum I 

Anterior uroneural neither 
fused with nor wedged 
into terminal vertebra 



Hiodontoidae 

(Fig- 4) 

IV 



Diagram i 



zool. 6, 6. 



Two postterminal centra, hypurals 
2 and 3 articulating with post- 
terminal centrum I 

Anterior uroneural neither fused 
with nor wedged into terminal 
vertebra. 



2I§§ 



332 



A CLASSIFICATION OF MODERN ISOSPOND YLOUS FISHES 



are considered to be basal members of a stock including the suborders Elopoidei, 
Clupeoidei, Gonorhynchoidei, and Stomiatoidei ; Salvelinus represents the Sal- 
monoidei ; Esox, the Esocoidei ; and Hiodon, the Osteoglossoidei and Mormyriformes. 
Of these four presumed lineages, the caudal skeletons of adult members of the 
Salmonoidei (II) are, with the exception of those of the Salangidae and of the neotenic 
aplochitonid Lovettia, the most easily recognizable. In this suborder the last few 
preterminal vertebrae have neural and hemal spines with flattened, anteroposteriorly 
oriented blades (Text-fig. 5). These together tend to make up a flange or keel running 




Fig. 3. Caudal skeleton of Esox lucius, x 2§. 



above and below the posteriormost portion of the vertebral column. This structure 
is developed to a variable degree in the different salmonoid fishes but is only totally 
absent in Lovettia and the salangids ; it is not found in the members of any of the other 
lineages, though Albula approaches it. In addition there is a series of about 10 
small, more or less s-shaped accessory (procurrent) rays in front of the caudal fin 
above and below in the salmonoids including Lovettia and the salangids ; these 
salmonoid accessory rays are easier to recognize than they are to differentiate from 
those that occur in many other isospondylous fishes. 

The osteoglossoid-mormyriform lineage (IV) has two quite different types of 
caudal skeleton, the one represented by Hiodon (Text-fig. 4) and the other by Osteo- 
glossum (Text-fig. 13), Heterotis (Text-fig. 14), and Mormyrus (Text-fig. 15). It 
would be difficult to confuse the second type with any other isospondylous caudal 



A CLASSIFICATION OF MODERN I SOS POND YLOUS FISHES 



333 



skeleton, but it is quite possible that Hiodon, by a different route, may have produced 
a fish with a caudal skeleton similar to that in the clupeoid stock. The main caudal 
feature militating against a hiodontid derivation for such a fish as Gonorhynchas 
is that this genus and the great majority of the other fishes here assigned to Lineage 
I have 17 branched caudal rays, whereas all of the members of the osteoglossoid- 
mormyriform lineage including Hiodon have 16 or fewer branched caudal rays. 

^NS of PR 
NS of TV 
NSotPT 

-HY8 
-HY7 
HY6 




Fig. 4. Caudal skeleton of Hiodon alosoides, x 4§ . In this fish the preterminal and 
terminal vertebrae and the first postterminal centrum each have a neural arch and spine 
that articulates with its centrum between the lateral uroneurals of the two sides. 



The haplomous lineage (III), as here understood, contains only the Esocidae, 
Dalliidae, and Umbridae. Chapman's (1944) assignment of the southern hemisphere 
aplochitonids, retropinnids, and galaxiids to the haplomous fishes is not accepted 
partly because of the caudal skeleton but also for other reasons that will be developed 
in Section 2 of this paper. The caudal skeletons of Esox (Text-fig. 3), Umbra and 
Novumbra are very similar and primitive. No caudal skeleton of Dallia has been 
available for investigation. 

The final lineage recognized here is that including the clupeoids and various 
other groups (I). About the only feature held in common by all its members is the 
high number (17 except in Coilia, Phractolaemus , and Cromeria) of branched rays 
in the caudal. This would seem to distinguish them from the osteoglossoid stock. 



34 



A CLASSIFICATION OF MODERN ISOSPONDYLOUS FISHES 



The absence of antero-posteriorly directed flanges on the last few neural and hemal 
spines would in turn separate them from the salmonoids. Nevertheless, Lineage 
I as accepted here may prove to be a heterogeneous assemblage. For example, 
such groups as the Gonorhynchoidei and Stomiatoidei may very possibly have 
arrived at caudal skeletons similar to those of the Clupeoidae by a process of parallel 
or convergent evolution rather than because of genetic relationship : about all 
that can be said for these questionable groups is that their caudal skeletons do not 
seem to show relationships with the salmonoid, esocoid, or osteoglossoid lineages. 




Fig. 5. Caudal skeleton of Hypomesus olidus, x iof . 



Suborder Elopoidei. Caudal skeletons of Elops, Megalops, Albula and Pterothrissus 
have been examined. All except the last have been discussed and figured by Hollister 
(1936, figs. 14-39). Though all are primitive, there is considerable difference between 
Elops and Megalops on the one hand and Albula and Pterothrissus on the other. 
In Elops and Megalops there are four pairs of uroneurals overlapping one another 
rather in the fashion of the body scales ; in the adult Albula and in Pterothrissus 
there are only two pairs of uroneurals and these are arranged almost linearly, one 
pair behind the other. In Elops and Megalops the terminal vertebra bears a neural 
arch and behind it lies a small median crest above and between the uroneurals of 
each side ; in Albula and Pterothrissus the terminal vertebra bears no neural arch 
and the median crest behind it has greatly expanded. 



A CLASSIFICATION OF MODERN ISOSPOND YLOUS FISHES 



335 



All of the species of Elopoidei appear to have 17 branched rays. Elops and Albula, 
but not Pterothrissus and Megalops, have a single bony fulcral plate at the front of 
the caudal base above and below. 

Superfamily Alepocephaloidae. The caudal skeleton of Alepocephalus (Text-fig. 
1) shows a peculiar combination of primitive (see above) and specialized features. 
There are the usual 17 branched caudal rays. Only one of the uroneurals is elongated, 
but this reaches forward to cover part of the first preterminal vertebra. Parts of 
two postterminal centra are visible, and hypurals 2 and 3 articulate with the first 
of these. Of specializations, the most notable perhaps is the long, curved, rod-like 
form of the upper epural and of the posterior neural spine. 




Fig. 6. Caudal skeleton of Chirocentrus dorab, x 4§ . 



Superfamily Clupeoidae. The caudal skeleton of Chirocentrus illustrated here 
(Text-fig. 6) differs from that of the other clupeids and engraulids examined (aside 
from Coilia) only in the somewhat enlarged and long-based neural arch on the 
terminal vertebra and the very slight development of a lateral spine on hypural 1 
(cf., Text-fig. 7). A specimen of Nematolosa examined, and illustrations of Clupea 
given by Ramanujam (1929, fig. 27) and of Anchoviella, Harengula, Opisthonema, 
and Sardinella by Hollister (1936, figs. 45-53) agree with Chirocentrus in all basic 
features. In these fishes the anterior uroneural is fused proximally with the terminal 
vertebra ; hypural 2 has no basal articulation ; one postterminal centrum is visible ; 
and the neural arch of the terminal vertebra has a dorsally projecting flange that 
is usually higher than broad. 

Many of these features have been developed in other isospondylous fishes but the 
loss of a basal articulation for hypural 2 apparently occurs only in portions of 
Lineage I. Normally, hypurals 2 and 3 articulate close together on a single centrum ; 



336 



A CLASSIFICATION OF MODERN ISOSPOND YLOUS FISHES 



if two postterminal centra are present hypurals 2 and 3 articulate with the first, 
which would consequently indicate that when two postterminal centra are present 
the first is of double origin. In the herring, however, Ramanujam (1929) has shown 
by embryological investigation that it is the terminal centrum that is of double 
origin (it seems to be formed by the fusion of the terminal vertebra with the front 
half of the first postterminal centrum), and that presumably with this change 
hypural 2 has lost its basal attachment. 

Though the anterior half of postterminal centrum 1 apparently always fuses 
into the terminal vertebra in the Clupeoidae, the fate of the more posterior centrum 
elements seems to vary. Thus, in Clupea, judging from Ramanujam's figures (1929, 
figs. 26, 27), the posterior half of postterminal centrum 1 fuses with the base of 
hypural 3 ; the same thing has probably occurred in Chirocentrus (Text-fig. 6) 



UN2 
I 




-HY3+PT 



Fig. 7. Terminal vertebra and associated structures of Dussumieria acuta, x 7J. 



and the majority of Clupeoidae. In Chirocentrus, therefore, the element labelled 
PT is probably postterminal centrum 2, fused with such centrum elements as may 
occur behind it. From Hollister's figures (1936, figs. 40-45) it would appear that in 
the dussumieriid J enkinsiella and perhaps in the engraulid Anchoviella, by contrast, 
the whole of postterminal centrum 1 fuses with the terminal vertebra. Finally, 
in Dussumieria it seems that the posterior half of postterminal centrum 1 has fused 
not only with the centrum elements behind it but with the base of hypural 3 as well. 
Dussumieria also differs from the other Clupeoidae examined in the fusion of hypural 
1 with the terminal vertebra and the anterior uroneural. 

Super families Chanoidae, Phractolaemoidae, and Knerioidae. The caudal fin 
of Chanos differs in no very essential way from that of the herring-like fishes. The 
uroneural is completely fused at its base with the terminal vertebra, but the first 
hypural is merely suturally articulated. In the single large specimen examined 
neither hypural 2 nor 3 has any connection with a centrum. The one significant 
difference from the herrings would appear to be that the median projection between 
the basal wings of the uroneurals is low, and just above it, extending along the 



A CLASSIFICATION OF MODERN ISOSPOND YLOUS FISHES 



337 



upper surface of the uroneural is a single long rod, presumably an epural. This 
structure is quite reminiscent of that in Alepocephalus (see Text-fig. i) though 
little else about the caudal skeleton is similar. 

The caudal skeletons of Phractolaemus and Kneria are in turn very similar to 
Chanos, differing from that of the herrings in approximately the same way as Chanos 
does. However, there are only 13 branched rays in the rounded caudal of Phracto- 
laemus. 

Superfamily Gonorhynchoidae. The caudal skeleton of Gonorhynchus (Text-fig. 8), 
as might be expected from the general consolidation of skeletal parts, shows a very 




Fig. 8. Caudal skeleton of Gonorhynchus greyi, x 7^. 



high degree of fusion for the isospondylous fishes. Not only are the uroneurals 
ankylosed to one another and to the terminal vertebra along with hypural 1, but 
hypurals 2 and 3 appear to have fused with one another. In general appearance 
the caudal skeleton of Gonorhynchus bears considerable resemblance to that of 
Dussumieria, but whether this is a result of convergent evolution or of genetic 
relationship remains in doubt. In any event, the caudal skeleton of Gonorhynchus 
gives no indication of relationship with the Osteoglossoidae, and its 17 branched 
rays would militate against a possible derivation from Hiodon. 

Suborder Stomiatoidei. Among the stomiatoid fishes the caudal skeletons of Pho- 
tichthys (Text-fig. 9) and Gonostoma were the only ones available. These two differ 
considerably from one another. In Photichthys neither the anterior uroneural nor 
hypural 1 is fused with the terminal vertebra, but in Gonostoma both appear to be. 
In both genera there are the usual 17 branched caudal rays. 



33» 



A CLASSIFICATION OF MODERN ISOSPONDYLOUS FISHES 



Superfamily Salmonoidae. The salmonids, thymallids, and coregonids (these 
terms are used here for groups of genera) all have a very characteristic and primitive 
caudal skeleton (Text-fig. 2). The anterior uroneural on either side bears a high 
crest between which the front ends of the epurals project. Two postterminal centra 
are visible ; hypurals 2 and 3 articulate with the first of these, but the base of hypural 
2 also extends somewhat forward on to the terminal vertebra. Salmo and Salvelinus 
have a well-developed neural spine on the first preterminal vertebra (Text-fig. 2), 
but in two skeletons of Thymallus (see also Lotz, 1864, pi. 10) and one of Coregonus 




Fig. 9. Terminal vertebra and associated structures of Photichthys argenteus, x j\ 



the epurals extend forward over but do not connect with the neural arch of the 
preterminal vertebra, thus replacing the neural spine in Salmo. 

Superfamily Argentinoidae. Among the Argentinoidae the only skeleton available 
is one of Argentina. This is here illustrated (Text-fig. 10) despite the strong possibility 
that the articulation between hypural 1 and the terminal vertebra is abnormal. 
Other available figures of the Argentinoidae are those of Opisthoproctus (Trewavas, 
x 933> pl- 2) and of Macropinna and Nansenia (Chapman, 19426, fig. 6 and 1948, 
fig. 12 respectively). All of these show the typical salmonoid neural and hemal spine 
expansions. They may, however, be separated from members of the Salmonoidae 
by the fact that the anterior uroneural never extends forward of the terminal vertebra 
and from the Osmeroidae by the presence of at least one postterminal centrum. 



A CLASSIFICATION OF MODERN I SOSPOND YLOUS FISHES 



339 



Subsequent examination of two stained specimens, 38 and 46 mm. in standard 
length, of the bathylagid Leuroglossus stilbius (kindly loaned by D. M. Cohen), 
shows the same basic structure described above. However, the salmonoid flanges 
are entirely lacking on the smaller and only slightly developed on the larger example. 
As compared with Argentina, there are, in Leuroglossus, only two uroneurals, which 
are very slender, and the anterior of these is fused with the terminal vertebra. 
The postterminal centrum is considerably smaller than that of Argentina (Text-fig. 
10), and only hypural 5 articulates with it. Also, Leuroglossus has no epurals. 




Fig. 10. 



Caudal skeleton of Argentina silus, x 6§. 



Super family Osmeroidae. Of the members of the Osmeroidae mentioned below, 
all but the Salangidae and Lovettia have the typical salmonoid neural and hemal 
spines (Text-fig. 5). None of them show a postterminal centrum or for that matter 
any other caudal resemblance to the esocoids. Though there is considerable variation 
among the members the most striking skeletal difference is that in Retropinna 
and Galaxias (Text-fig. 11) the anterior uroneural is not fused with the terminal 
vertebra whereas in Hypomesus (Text-fig. 5), Plecoglossus (Text-fig. 12), Aplochiton, 
and the salangid Leucosoma it is. The caudal skeleton of the last-named genus is 
quite different from all of the others. 

In one respect the caudal fins of the southern Osmeroidae differ from those of 
the northern members of the group (also from the Argentinoidae and Salmonoidae). 
All of the latter, including the salangids and Plecoglossus, have 17 branched caudal 



zool. 6, 6. 



2I§§§ 



34° 



A C 



CLASSIFICATION OF MODERN ISOSPOND YLOUS FISHES 




Fig. ii. Caudal skeleton of Galaxias fasciatus, x 6f. 

V HY7 




Fig. 12. Caudal skeleton of Plecoglossus altivelis, x 8. 



A CLASSIFICATION OF MODERN ISOSPOND YLOUS FISHES 



34* 



rays. In the southern Retropinna and Prototroctes there are 16 branched caudal 
rays ; and in Aplochiton, Lovettia, and Galaxias 14. 

Suborder Esocoidei. The caudal skeletons of Esox (Text-fig. 3) and of Umbra 
are quite similar. That of Umbra appears the more primitive of the two, with six 
" hypurals " of about equal size between a narrower strut above and below. The 
upper of these struts is the uppermost hypural ; the lowermost is the hemal arch 
from the second preterminal vertebra, the first preterminal vertebra giving rise to 
one of the broad " hypurals ". In seven specimens of Esox lucius for which caudal 
rays were counted, five had 17 branched rays and two 16 ; for three specimens 
of Umbra krameri, two had 9 branched rays and one 8. 




Fig. 13. Caudal skeleton of Osteoglossum bicirrhosum, x 5 J. 



Superfamily Hiodontoidae. As previously mentioned, one of the most primitive 
of isospondylous caudal skeletons is certainly that of Hiodon. In two skeletons 
examined, one of H. tergisus and one of H. alosoides, considerable variation appeared, 
not only from fish to fish, but on the two sides of the same fish. Thus the skeleton 
of H. alosoides has three uroneurals on one side (Text-fig. 4) and four on the other. 
In H. tergisus both the terminal and the first postterminal centra have double 
neural arches, and the anterior tip of the single epural runs in between but is free 
from the sides of the posteriormost arch. In H. alosoides (Text-fig. 4) both the terminal 
and the first postterminal vertebrae have single neural arches with spines that 
nearly reach the caudal fin base. Nine specimens of H. alosoides and one of H. tergisus 
examined all had 16 branched caudal rays. 

Super families Osteoglossoidae and Notopteroidae. The caudal skeletons of Osteo- 
glossum (Text-fig. 13), Pantodon, and Heterotis (Text-fig. 14), while showing strong 



342 



A CLASSIFICATION OF MODERN ISOSPOND YLOUS FISHES 



similarities to one another, are so different from those of all the foregoing isospondy- 
lous fishes as to be somewhat difficult to interpret. (Notopterus, see Whitehouse, 
1910b, pi. 47, fig. 6 is even more specialized in the same direction, and a skeleton 
of Arapaima was not available.) Presumably, the terminal vertebra is that preceding 




Fig. 14. Part of caudal skeleton of Heterotis niloticus, x 4§ . 




Fig. 15. Caudal skeleton of Mormyrus cdballus, x 5 J. 



the one to which two hypurals are attached. If this is correct, postterminal centrum 
1 is present in an unusually complete form, and postterminal centrum 2 has fused 
with the large upper hypural plate and with the uroneurals as well (the latter fusion 
is incomplete in Heterotis, Text-fig. 14). Though any other interpretation of these 
caudal skeletons raises even more difficulties, the one just given presents three : 
first, hypural 3 in Osteoglossum has lateral flanges that are not found elsewhere in 



A CLASSIFICATION OF MODERN I SOSPOND YLOUS FISHES 343 

isospondylous fishes ; second, postterminal centrum 1 in Osteoglossum has a neural 
spine instead of the usual epural structure ; finally, and most oddly, the uroneurals 
in all three genera under consideration fuse with . postterminal centrum 2 instead 
of with the terminal vertebra. Because of this last peculiarity, the alternate inter- 
pretation that what is here called postterminal centrum 2 is really the terminal 
vertebra deserves serious consideration. In any event, it is difficult to see how the 
caudal structure of these fishes has been derived from any of the isospondylous 
groups, including Hiodon, dealt with up to here. Heterotis has only 12 branched 
caudal rays, and Osteoglossum 14. 

Order Mormyriformes. The caudal skeleton of Mormyrus caballus (Text-fig. 15) 
is basically very similar to those of Heterotis and Osteoglossum. The main differences 
lie in the flange on hypural 2 and the number of caudal rays in Mormyrus (16 
branched) . 

DISCUSSION 

Two subjects call for discussion. One is the functional significance of the different 
types of caudal skeletons noted above. The other concerns the bearing of the caudal 
structure on the limits of the order Clupeiformes. 

Two aspects of the relation between structure and function will be considered 1 . 
The first of these concerns the elongate uroneurals of the primitive isospondylous 
fishes. According to Regan (1910a) and Ramanujam (1929) the uroneurals are, 
at least primarily, modified portions of neural arches of postterminal centra. What- 
ever their derivation, the elongation of these uroneurals in the so-called primitive 
isospondylous caudal skeleton is in itself a specialization. The only obvious explana- 
tion for these long uroneurals is that they stiffen the upturned terminal portion 
of the vertebral column. With the more abrupt upturning in this area found in 
advanced members of both the clupeoid and salmonoid lineages, the uroneurals, 
which start out as one or more shields along the sides of the postterminal centra, 
develop into struts which are at first wedged into the terminal vertebra and in 
more " advanced " forms fuse with it. Apparently a short, anchored strut serves 
here better than a long cover. 

The second functional aspects has to do with the relationship between the caudal 
skeleton and the shape of the tail. In a very broad sense there is a correlation between 
these two features. Very generally, when the tail is forked the caudal skeleton is 
divided into distinct upper and lower portions, but when the tail is rounded the 
posterior outline of the hypurals is rounded, often with a number of hypurals of 
subequal size (e.g., Amia, Umbra) or with a central one somewhat larger than the 
others as in the cods. However, this relationship is far from strict. For example, 
the round-tailed serranid Epinephelus has the caudal skeleton of a typical fork- 

1 In this discussion the caudal skeleton is treated as if it were an independent functional unit. Ob- 
viously it is not, for beside providing an axis for the basal articulation of the caudal rays it serves as a 
source of attachment for some of the caudal musculature (cf. Hindersson, 1910 ; Schmalhausen, 
especially 191 3 ; and Greene and Greene, 19 14) and for the transmission of the blood vessels and nerves 
to the caudal fin. However the role that these various functions have played in the evolution of the caudal 
skeleton are almost entirely unknown at the present time and have consequently been left out of con- 
sideration. 



344 A CLASSIFICATION OF MODERN ISOSPOND YLOUS FISHES 

tailed fish. Conversely, the fork-tailed Mormyrus has a caudal skeleton that should 
belong to a round-tailed fish. Apparently the shape of the tail takes a very long 
time to imprint itself on the structure of the caudal skeleton. 

Similarly there seems to be a relationship, though again not a strict one, between 
the shape of the caudal fin and the number of caudal rays. In fork-tailed fishes the 
number is usually constant within groups. Indeed the basic number of 17 branched 
caudal rays runs through two of the four isospondylous lineages postulated here. 
However, a rounded caudal is often associated with a lower (and variable) number 
of principal caudal rays. Presumably, in a fork-tailed fish the greatest strength 
and functional importance lies in the outermost principal caudal rays which form 
rigid limits to the fin. In round-tailed fishes these outer rays lose their significance 
and the fin tapers from the middle to above and below. One is inclined to wonder 
whether those fork-tailed isospondylous fishes with some lower caudal count than 
the typical one, e.g., Hiodon, Aplochiton, have not had some round-tailed ancestors? 

With regard to the information caudal structure can provide on the limits of 
the order Clupeiformes, there are only three groups that need be considered here : 
the haplomous, mormyriform, and iniomous fishes. Each of these has been included 
in or excluded from the Clupeiformes, depending on the classification adopted. 

The first two of these groups have already been dealt with. Suffice it to say here 
that so far as the caudal skeleton is concerned the haplomous fishes appear to be as 
primitive as any of the isospondylous fishes. Since they stand at the base it is impos- 
sible to derive them from any other modern isospondylous fish, so far as the caudal 
skeleton is concerned. 

The mormyroid caudal skeleton seems clearly to have been developed from the 
type now found in Heterotis. The structural inference, as already noted, is that the 
mormyrids retain not only the imprint established in a round-tailed form, but one 
of a peculiar and definitely identifiable type. 

Among the iniomous fishes (Scopeliformes) only skeletons of Neoscopelus, Lam- 
panyctus, Aulopus, and synodontid fishes have been available for examination. 
The caudal structure of these seems to be somewhat more primitive than that of 
many isospondylous fishes in that one postterminal centrum is clearly visible. In 
certain features, e.g., the flanged uroneural, these iniomous caudal skeletons appear 
to approach the salmonoid Clupeiformes ; however, they lack the expanded posterior 
neural and haemal spines of the salmonoids. In sum, there is little beyond an indica- 
tion of possible relationships here. 

Whether any or all of the three groups discussed above should be included in or 
excluded from the order Clupeiformes will, of course, depend in the first place on 
the nature of the whole animals, not just their tails, and in the second on a consensus 
of human opinion regarding what constitutes a fish order. 



SECTION 2. THE CLASSIFICATION AND RELATIONSHIPS OF THE 

SALMONOID FISHES 

The members of the Suborder Salmonoidei (as defined here) today are the dominant 
fishes in the coldest fresh water of both hemispheres. They are also represented in 



A CLASSIFICATION OF MODERN ISOSPOND YLOUS FISHES 345 

the temperate and arctic regions of northern seas by one group (Osmeridae) and in 
deeper oceanic waters by another (Argentinoidae 1 ) . 

It is hardly surprising that there should be difficulty in defining as large and varied 
an assemblage as the salmonoids. Regan (1929) and others have differentiated them 
from the clupeoid groups by the absence or incompleteness of the oviducts. However, 
Kendall (1921) demonstrated that the oviducts of Salmo and Osmerus " are not 
radically different from those of other isospondylous fishes ". Regan (1913^ : 289) 
gives the following additional salmonoid characters : an adipose fin usually present, 
one supramaxillary or none, and parietals well developed. Probably the most 
useful of these three features for purposes of differentiation is the adipose fin, for 
it is found elsewhere in the isospondylous fishes only among the stomiatoids and 
the Rosauridae. However, there are salmonoids (Galaxias) and salmonoid derivatives 
(the haplomous fishes) in which the dorsal fin has moved far back on the body and 
presumably squeezed out the adipose fin (Garstang, 1931 : 246-250). An adipose 
is also lacking in at least one of the oceanic salmonoids (Microstoma) . An additional 
diagnostic trait of most salmonoids is that the last few "preterminal" vertebrae 
have laterally compressed neural and hemal arches, forming a flange above and 
below the posterior portion of the vertebral column (see previous section). This 
flange seems to be lacking only in the neotenic Lovettia and the Salangidae among 
the salmonoid skeletons examined. At least the posteriormost of these neural and 
hemal spines lead out to a series (usually about 10) of characteristic small, curved, 
accessory rays ; these occur prominently in both Lovettia and the salangids. A 
final trait that will apparently distinguish the salmonoids (and haplomous fishes) 
from the herrings and their relatives is the absence of a lower series of intermuscular 
bones, i.e., the series that articulates proximally with the base of the lower ribs 
(cf., Emelianov, 1935.) 

The present classification of the salmonoid families stems from Regan (1913a : 
289). There, he differentiates the families as follows : 

"I. An orbitosphenoid ; an opisthotic ; a mesocoracoid ; vertebrae upturned at base 

of caudal fin . . . . . . . . .1. Salmonidae 

II. An orbitosphenoid; no opisthotic; no upturned vertebrae; mesopterygoids toothless. 

A mesocoracoid ; parapophyses inferior . . . . . 2. Argentintdae 

No mesocoracoid ; parapophyses lateral . . . . 3. Microstomidae 

III. No orbitosphenoid ; no opisthotic ; no upturned vertebrae ; mesopterygoid 
toothed (absent in the Salangidae). 

A. A mesocoracoid ; maxillaries dentigerous, entering gape . . 4. Osmeridae 

B. No mesocoracoid ; maxillaries dentigerous, entering gape. 

Head compressed ; mesopterygoid well-developed dentigerous ; ribs ossified 

5. Retropinnatidae 
Head strongly depressed ; no mesopterygoid ; ribs not ossified . 6. Salangidae 

C. No mesocoracoid ; maxillaries toothless, behind praemaxillaries. 
Praemaxillaries not extending whole length of maxillaries ; roof of myodome 

unossified ; no adipose fin. ....... 7. Galaxiidae 

Praemaxillaries nearly reaching extremities of maxillaries ; roof of myodome 

ossified ; an adipose fin ...... 8. Haplochitonidae" 

1 The name Argentinoidae, rather than Opisthoproctoidae, will be used throughout this paper for the 
perhaps over-simple reason that the most recent workers on the group (Hubbs, 1955 ; Bertelsen, 
1958 ; and Cohen, 1958) seem to prefer the former name. 



346 A CLASSIFICATION OF MODERN ISOSPOND YLOUS FISHES 

With regard to this classification, Regan goes on to say (op. cit.: 290) : 

"It is of some interest to note that the Galaxiidae and Haplochitonidae are 
related to, but more specialized than, the Osmeridae, or Smelt family, of northern 
seas. Retropinna, from the coasts and rivers of Australia and New Zealand, is still 
nearer to the Galaxiidae and Haplochitonidae." 

Subsequent work by Trewavas (1933), Parr (1937), and Chapman (19420, 19426, 
1943, and 1948) has abundantly shown that the members of Regan's section II — 
including his Argentinidae and Microstomidae — are rather widely divergent from 
the other salmonoid fishes. Characteristic features of the group are the toothless 
upper jaw, the pelagic eggs, etc. The classification of the families within the group 
seems to be rather controversial (cf., the above papers, Hubbs, 1953, and Bertelsen, 
1958) but the subject will not be entered upon here. 

Similarly the members of Regan's (1913a) section I may be readily distinguished 
from all other members of the suborder by the three upturned caudal vertebrae. 
(Other members of the suborder have one or, at most, two.) Whether the family 
Salmonidae should be interpreted in the broad sense of Regan (1913a) or whether 
the Thymallidae and /or Coregonidae should be split off as separate families are 
matters that can best be left to those more familiar with these fishes to decide. 
However, the assignment of the fossil Thaumaturidae by Voigt (1934), Berg (1940) 
and others to the salmonid group seems to be open to some question. The typical 
salmonoid flanges formed by the last few neural and hemal spines are lacking, as 
is the adipose fin (ace. Voigt) ; the upward inflexion of the last three vertebrae 
shown in Voigt 's pi. 2, fig. 5 is not salmon-like and is, in any event, not duplicated 
in the specimens shown in pi. 1, figs. 1 and 2 ; and the posterior position of the dorsal 
is more reminiscent of the haplomous fishes than of the salmons. 

The last group of salmonoid fishes — section III of Regan — contains a rather 
diverse assemblage of fishes. It can, however, be immediately divided into a northern 
group of families — Osmeridae, Plecoglossidae, and Salangidae — with 17 branched 
caudal rays, and a southern hemisphere group — Retropinnidae, Aplochitonidae, 
and Galaxiidae — with fewer branched caudal rays. This subdivision is in line with 
Regan's (1913a) statements quoted earlier. Nevertheless both groups are quite 
varied within themselves, and each has given rise to a series of specializations partly 
paralleled within the other. Thus the northern Plecoglossus with its specialized 
dentition to some extent resembles the southern Prototroctes with its small flexible 
teeth, and the northern neotenic salangids are partly duplicated in the southern 
neotenic Lovettia. 

The salmonoid allocation of the three southern hemisphere families remained 
uncontested until Chapman (1944), on the basis of an osteological study of Aplochiton, 
transferred the Aplochitonidae, Galaxiidae, " and very probably the Retropinnidae " 
to the haplomous fishes. Subsequent authors, e.g., Schultz and Stern (1948) and 
Wilimovsky (195 1), have generally accepted Chapman's allocation. However, 
Blackburn (1950), from an investigation of the biology of the aplochitonid Lovettia 
seali, points out " that the life history of this species is similar to that of various 
salmonids, which makes the author incline to the views of Regan and Jordan " 
with regard to its systematic position. 



A CLASSIFICATION OF MODERN ISOSPOND YLOUS FISHES 347 

In view of this difference of opinion, the British Museum material of the southern 
hemisphere forms has been examined. A general review of these, at least insofar 
as their systematic position is concerned, is undertaken in the following paragraphs. 

In general the three southern families can be divided into two groups — the anadro- 
mous retropinnids and aplochitonids {Prototroctes!) with an adipose fin and a forked 
caudal, and the fresh-water or catadromous galaxiids without an adipose fin and 
with a rounded caudal. 

The galaxiids would appear to be a fairly homogeneous group even though they 
are represented in southern South America, southern Africa, New Zealand, Tasmania, 
Australia, and presumably New Caledonia. Regan (1905) has an early revision of 
the family ; more recently Scott (1936, etc.) and Stokell (1945, etc.) have worked 
on the systematics of the group. Swinnerton (1903) has written on the osteology of 
one of the species. 

By contrast with the galaxiids, the retropinnids and aplochitonids are most 
diverse. The Retropinnidae has only a single Australian and New Zealand genus 
revised by Stokell (1941). The Aplochitonidae contains three very different genera : 
a fairly normal Aplochiton from southern South America and the Falkland Islands, 
Proctotroctes with a highly specialized dentition and with two almost if not entirely 
extinct species from Australia, Tasmania, and New Zealand, and the neotenic 
Lovettia from Tasmania. As already noted, Chapman (1944) has a paper on the 
osteology of Aplochiton, and Blackburn (1950) has published on the biology of 
Lovettia. A general account of the New Zealand forms of all three families is to be 
found in Stokell (1955). 

In external characters, aside from those already mentioned, Retropinna and 
Proctotroctes are scaled ; the other southern genera are naked. Retropinna has 
somewhat the look of, and is called, smelt (Stokell, 1955 : 9-18, pis. 4 and 5) whereas 
Prototroctes looks rather like a Coregonus (Stokell, 1955 : 41-44, pi. 13). Aplochiton 
is more trout-like (Eigenmann, 1927 : pi. 15) ; Lovettia has a definitely larval 
appearance (Blackburn, 1950, pis. 1 and 2) ; and the galaxiids (Stokell, 1955 : 
19-40, pis. 6-12) have somewhat the look of the northern Umbra. 

So far as I can see the rather variable fin counts have no significance for the classi- 
fication of the group as a whole except for the caudal. Judging from counts in a 
single specimen of each genus, Prototroctes and Retropinna have 16 branched rays, 
whereas Aplochiton, Lovettia and Galaxias have 14 branched. All of these genera 
have the typical salmonoid accessory rays (see previous Section) at the front of the 
caudal above and below. 

The dentition of these southern hemisphere forms is most varied. At the one 
extreme are certain species of Retropinna (Stokell, 1941) in which there is a long 
posterior portion of the maxillary, a short premaxillary, a crescent-shaped vomer, 
palatines, mesopterygoid, tongue, a long bone covering the basibranchials, pharyn- 
geals, and dentary all with large teeth. At the other extreme is Prototroctes with 
fine, peculiarly-shaped, flexible teeth in the upper jaw and apparently none in the 
lower. 

The maxillary is also variable. In the single genus Retropinna it may be toothed 
or toothless (Stokell, 1941 : pi. 55). In the three genera of the family Aplochitonidae 



348 A CLASSIFICATION OF MODERN ISOSPOND YLOUS FISHES 

the toothless maxillary is excluded from the gape as effectually as it is in the perch- 
like fishes. A supramaxillary seems to be consistently absent. 

In Galaxias and Lovettia the premaxillaries appear to be slightly protractile ; 
in both these genera they have rudimentary anterior pedicels. 

With regard to the skull the parietals meet on the midline except in Lovettia. 
In a ripe female specimen of Lovettia both the frontals and parietals are elongate 
bones running along the superolateral borders of the skull. The whole median area 
of the skull roof is merely membranous, with the brain clearly visible. 

The anterior ends of the frontals project well forward over the ethmoid, not only 
in Lovettia but in all the other southern genera as well. There are no proethmoids 
in any of them and what Chapman describes as proethmoids in Aplochiton (1944 : 
150, fig. 1) are not separate elements in British Museum specimens but are merely 
anterior prongs of the frontals. (If these anterior portions somehow came to develop 
as separate elements in the northern haplomous fishes, would they not be proeth- 
moids?) 

Whether the mesethmoid is cartilaginous or bony in Lovettia is impossible to 
say ; it is certainly an endochondral formation. However, in Galaxias (cf., Swinner- 
ton, 1903) there is a dermal mesethmoid as well. 

I have not seen an orbit osphenoid in any of the southern hemisphere forms. 

At the back of the skull the small supraoccipital is excluded from the foramen 
magnum in all the southern genera. 

In the suspensorium the large mesopterygoid is usually toothed in southern 
forms. The metapterygoid extends forward over much but not all of the quadrate. 
There seems to be no knob behind the palatine for articulation with the lateral 
ethmoid (prefrontal). As to the ectopterygoid, it and the palatine seem to be fused 
into a single long, mostly toothed bone in Retropinna. However in Prototroctes 
the two elements are separate. 

The suboperculum is a rather large bone forming part of the gill cover in all except 
Lovettia. In Lovettia, the suboperculum gives the distinct impression of being the 
seventh (and terminal) branchiostegal. 

In the pectoral girdle, none of the southern forms has a mesocoracoid. 

In the caudal skeleton there are never any visible " postterminal centra " (see 
previous section). The anterior uroneural may (Galaxias) or may not (Prototroctes) 
be fused with the "terminal vertebra". In all, except Lovettia, the salmonoid 
flattening of the last few neural and hemal arches is distinctly developed. 

As to soft anatomy, oviducts appear to be lacking in the females. In adult males 
of Lovettia, which seems to be unique in this respect, the urinogenital openings move 
forward to just behind the pectorals. None of the southern genera have pyloric 
caeca. Lovettia has a closed air bladder quite unlike that of the osmerids, salmo- 
nids, or northern haplomous fishes including salangids (N. B. Marshall, personal 
communication) . 

With this background regarding the southern hemisphere Aplochitonidae, Retro- 
pinnidae, and Galaxiidae, it remains to compare them with the northern hemisphere 
Esocoidei and Osmeroidae. 

The northern haplomous fishes have been defined by Chapman (1934 : 372) as 



A CLASSIFICATION OF MODERN ISOSPOND YLOUS FISHES 349 

follows (and the definition has been accepted with little change by Berg, 1936, 
1940, and 1955) : 

" Physostomous teleosts with the pelvic fins abdominal ; cycloid scales ; paired 
proethmoids ; parietals separated by the supraoccipital ; maxillary forming part 
of the edge of the upper jaw but lacking teeth ; premaxillary not protractile ; 
head scaly ; the dorsal and anal fins far back on the body ; but without adipose 
fin, mesocoracoid, or orbitosphenoid, and with no true spines in the fins." 

As has already been noted, the southern hemisphere Lovettia has a closed air 
bladder ; none of the southern forms have paired proethmoids ; only Lovettia 
has the parietals in any degree separated by the supraoccipital ; the maxillary is 
excluded from the gape in the Aplochitonidae, and in some species of the Retro- 
pinnidae the maxillary is toothed ; in Lovettia and Galaxias the premaxillary appears 
to be slightly protractile ; none of the southern forms have scales on the head ; 
the dorsal in Proctotroctes is not especially far back ; and only the galaxiids lack 
an adipose. 

If, then, the southern hemisphere families are included in the Esocoidei, only 
the following portions of Chapman's (1934) definition are left intact : 

" Teleosts with the pelvic fins abdominal ; cycloid scales ; anal fin far back 
on the body ; without mesocoracoid or orbitosphenoid, and with no true spines in 
the fins." 

The above words would apply, except for the absence of a mesocoracoid, to most 
Osmeroidae, and in toto to the included family Salangidae. 

This is not to say that the southern families do not have haplomous resemblances, 
for they do. Indeed, some, but by no means all, of the similarities between Aplochiton 
and the haplomous fishes which Chapman lists (1944 : 164) will hold for all three 
southern families. Two more may be added here. First the southern families, 
like the northern Esocoidei, lack pyloric caeca, whereas the northern osmeroids 
(except the Salangidae) have several to very numerous caeca. Second, the southern 
forms all have fewer than 17 branched caudal rays, whereas the northern osmeroids 
all have 17 branched rays ; in the northern haplomous fishes there is a reduction 
in caudal ray number except in Esox, and even here two of the seven specimens 
examined had fewer than 17 branched rays. 

In comparing the southern families with the northern Osmeroidae, Chapman 
(1944 : 163) has already noted a number of differences, and several more may be 
added. The most important would seem to be the following : in the north a supra- 
maxillary, mesocoracoid, epipleural ribs, and pyloric caeca are present (except 
in the Salangidae), whereas in the southern forms they are absent ; and in the northern 
forms there are 17 branched caudal rays whereas there are 16 or fewer in the southern 
families. Admittedly these differences in the southern group are all modifications 
away from the basic salmonid type and in a haplomous direction. The same may 
be said of the posterior dorsal and anal position and the loss of an adipose in the 
galaxiids, which, of all the southern forms, have progressed farthest along this 
route. However, this direction has also been followed to a considerable extent by 
the northern salangids. 

If, however, it is admitted that all three southern families are related and if the 



35o 



A CLASSIFICATION OF MODERN ISOSPOND YLOUS FISHES 



most primitive features of these families are added together, we have a fish that is 
basically osmerid. In Prototroctes the dorsal is median and the adipose is well 
developed. In some species of Retropinna the maxillary is toothed and indeed the 
whole dentition is osmerid, with the teeth of the tongue and basibranchials biting 
between those of the palatines and mesopterygoids. At the other end of the fish 
the salmonoid flanges on the neural and hemal spines are present (except in Lovettia) 

OSMERIDAE and 

PLECOGLOSSIDAE 

proethmoids sometimes 

present 

SALANGIDAE 
no pyloric caeca 
no mesocoracoid 
no flanges on neural and 

hemal spines APLOCHITONIDAE 

v RETROPINNIDAE 

and GALAXIIDAE 
6 or fewer branched 
caudal rays 
no pyloric caeca 
no mesocoracoid 
adipose fin present or 

absent 
maxillary toothed or 
toothless 




SALMONOIDAE 
17 branched caudal rays 
no proethmoids 
pyloric caeca present 
mesocoracoid present 
an adipose fin 
a toothed maxillary 
flanges from posterior neural and 
hemal spines 



ESOCOIDEI 
usually fewer than 17 branched 

caudal rays 
proethmoids present 
no pyloric caeca 
no mesocoracoid 
no adipose fin 
no toothed maxillary 
no flanges from posterior neural 

and hemal spines 




Diagram 2 
along with the curved accessory caudal rays, and the caudal supporting skeleton 
is osmerid (see Section 1). 

This last feature seems particularly important in view of the fact that the caudal 
skeleton of the haplomous fishes is not only very different but far more primitive 
than that of the northern and southern Osmeroidae. Finally, the anadromous life 
history of several of the southern forms recalls that of many salmonids. From all 
this it seems to the present author that the Aplochitonidae, Retropinnidae, and Gal- 
axiidae are derivatives of a proto-osmerid stock that have evolved, some more and 
some less, in a haplomous direction ; and that this same direction has been followed 
a third time to a lesser extent by the Salangidae (see accompanying diagram). 



A CLASSIFICATION OF MODERN ISOSPOND YLOUS FISHES 351 

That the Salangidae and the southern hemisphere Osmeroidae have departed in 
the same direction from the northern Osmeroidae is indicated by the characters 
listed in the diagram. But the similarities between the salangids and the aplochitonid 
Lovettia are even more striking. Both Lovettia and the salangids are more or less 
neotenic, anadromous fishes, with tremendous sexual differentiation. The sexual 
differentiation, however, is of a very different type in the two groups : in the salangid 
males, but not the females, there is a series of large scales above the anal fin, and 
the fin itself has a peculiar curvature in several of the median rays ; in the adult 
male Lovettia the urinogenital papilla is just behind the pectoral fins. In the head 
of both, the posterior portion of the skull is very incompletely roofed by the frontals 
and parietals. Furthermore, these are the only two groups in the Salmonoidei 
that have no flanges on the preterminal neural and hemal spines. 

In view of this it may appear that Lovettia and the salangids are closely related. 
Again I would prefer the hypothesis of parallel (neotenic) development. That 
Lovettia belongs with the southern osmeroids, and the salangids with the northern 
forms is indicated by the caudal ray count. In dentition, the salangids have a toothed 
maxillary typical of northern osmeroids, whereas Lovettia has a toothless maxillary 
excluded from the gape typical of the other two genera of aplochitonids. Additional 
characters for differentiating at least Salangichthys microdon from Lovettia sealei 
are the presence of a high (cartilaginous?) lump just behind the palatine for articula- 
tion with the lateral ethmoid (prefrontal) in Salangichthys and the absence of such 
a structure in Lovettia ; the presence of a well-developed subopercle in Salangichthys 
whereas the subopercle resembles a free branchiostegal in Lovettia ; the six branchio- 
stegals of Lovettia vs. the four in Salangichthys ; and finally the normal, open sal- 
monoid air bladder in Salangichthys vs. the peculiar closed one in Lovettia (see above). 

A final question regards the interrelationships of the members of the southern 
group. Berg (1940) separated the galaxiids as a distinct order, primarily on the basis 
of brain structure. Unfortunately, all that is known about the brain of these southern 
forms seems to be a single remark by Swinnerton (1903) regarding Galaxias. However, 
there are other features that set Galaxias apart from the other southern osmeroids. 
Among these may be mentioned the general body form and the catadromous habits. 

Another line of differentiation within the southern section of the Osmeroidae 
is shown by Lovettia with its neoteny, sexual differentiation, and closed swimbladder. 
Such peculiarities would certainly warrant family differentiation from the aplochi- 
tonids, except that the Aplochitonidae is already a family of only three genera. 
Furthermore, the two remaining genera — Prototroctes and Aplochiton — are also 
very different from one another, and it seems preferable not to erect three separate 
families each with a single genus. 

Nevertheless, the fact remains that the southern Osmeroidae are now represented 
by several highly differentiated forms. Indeed, the differentiation is as great as in 
the northern Osmeroidae. 

So far as the haplomous fishes are concerned, Berg stated in 1936 that the " Eso- 
coidei are a specialized group of Clupeiformes originating from Osmeroid fishes at 
the end of the Cretaceous ". The present paper would tend, like that on Aplochiton 
by Chapman (1944), to demonstrate an even closer set of morphological links between 



352 A CLASSIFICATION OF MODERN ISOSPOND YLOUS FISHES 

osmeroids and esocoids than was known to Berg. However, the primitive caudal 
skeleton of the haplomous fishes indicates (see previous section) that Esox, Umbra, 
etc. have been derived from a somewhat less advanced salmonoid stock than the 
northern and southern Osmeroidae. 

From the foregoing account, a classification (to superfamily) of the modern 
salmonoid fishes and their haplomous derivatives may be written as follows : 

Suborder Salmonoidei 

Superfamily Salmonoidae 

Superfamily Argentinoidae (= Opisthoproctoidae) 

Superfamily Osmeroidae 

Suborder Esocoidei 

Superfamily Dallioidae 
Suprfamily Umbroidae 
Superfamily Esocoidae 

In this classification the three haplomous superfamilies of Chapman (1934) and 
Berg (1940, 1955) have been accepted without any attempt at critical evaluation. 
As to the salmonoid superfamilies listed here, everyone admits that they are related ; 
the only question is how to express the relationships. Here, the salmonid and osmerid 
groups have been recognized as separate superfamilies, and the argentinoids have 
been demoted to a superfamily equivalent to them. The first action is taken because 
it seems advisable, where feasible, to segregate a basal stock that has given rise 
to a number of lineages from the advanced members of those lineages. 

The place that the Argentinoidae should hold in relation to the other two salmonoid 
superfamilies is somewhat less clear. Nevertheless it can only obscure the relation- 
ship that undoubtedly does occur to set the Argentinoidae up as a suborder (Opistho- 
proctoidei of Berg, 1940, Chapman, 1942^, etc.) with a taxonomic rank equivalent 
to that of the salmonoids themselves. In the specializations of the head skeleton 
(Chapman, 1942a and b), in the loss of a ventral mesentery (Kendall, 1935, Table I), 
and in the caudal skeleton (see previous section), the Argentinoidae is more speci- 
alized than the Salmonidae 1 . It would seem that the Argentinoidae is a much more 
divergent offshoot of the stock represented by Salmo today than is the Osmeroidae. 
But to express this greater divergence in a classification seems to lead to more 
incongruities than to consider them as a superfamily equivalent to the Salmonoidae 
and Osmeroidae. 

SECTION 3. THE GONORHYNCHOID FISHES AND THEIR 

RELATIONSHIPS 

The single modern genus generally attributed to the suborder Gonorhynchoidei 
is Gonorhynchus (cf., Regan, 1929 ; Berg, 1940 ; Wilimovsky, 1951 ; etc.). The 
systematic position of the suborder has always been dubious. Ridewood (19056), 
on the basis of a study of the skull of Gonorhynchus greyi, while emphasizing its 
isolated position, tentatively suggests a distant relationship with the salmonids. 

1 Another distinguishing character usually attributed to the argentinoids is the presence of a spiral 
valve in the intestine (cf., Kendall & Crawford, 1922). However, Cohen (1958 : 97, 98) has recently 
demonstrated that the " spiral valve " of argentinoids is not so diagnostic of that group as has usually 
been stated. 



A CLASSIFICATION OF MODERN I SOSPOND YLOUS FISHES 353 

Regan (1929), in his synopsis of the isospondylous fishes, places the gonorhychids 
at the end of the order, behind the mormyrids. His basis for this placement is 
that the gonorhynchids have the " parapophyses ankylosed with centra, appearing 
as strong processes". Subsequent authors, e.g., Berg (1940), Wilimovsky (1951), 
have generally accepted Regan's allocation. 

That the parapophyses in Gonorhynchus are strong processes fused with the centra 
is true, but the value of this character in the classification of the isospondylous 
fishes seems dubious. For example, among the groups included by Regan in the 
section with Gonorhynchus, this genus and Osteoglossum have " strong processes " 
but Hiodon and Mormyrus appear to have none whatever, the ribs articulating 
directly with the centra. 

A search for other characters that might indicate the relationships of Gonorhynchus 
discloses the following, all of which point toward an albulid-clupeid relationship. 
There are 17 branched caudal rays (this could equally well indicate a salmonoid 
ancestry). The caudal skeleton, except for the greater fusion of parts, resembles 
that of Dussumieria, the upright neural arch on the terminal vertebra being especially 
reminiscent of the clupeoids. Arising from the parapophyses are two sets of seg- 
mental bones ; one of these undoubtedly represents the lower ribs, but the other 
seems to be the lower intermuscular bones. Lower intermuscular bones again are 
characteristic of the clupeoids and seem to be absent from most, if not all, other 
isospondylous groups. 

As to relationships within the great clupeoid stock, it is here suggested that the 
gonorhynchids belong with the chanids, phractolaemids, kneriids, and cromeriids. 
These five groups are so widely different that any relationship between them is 
difficult to comprehend. Yet the following similarities may be marshalled. 

In external characters, the gill membranes are always attached to the isthmus, 
and in the three African families the gill openings are considerably restricted. In 
all, the mouth is small and toothless, or nearly so. The preopercular border is 
free only below (Chanos), if at all. 

In the head the maxillaries run forward behind the premaxillaries and nearly 
or quite meet on the midline. There are no supramaxillaries. A preorbital bone 
extends forward above the upper jaw nearly to the end of the snout (this is least 
developed in Chanos). The preopercle extends only a short way up the side of the 
head and there may be a suprapre opercular (Phractolaemus and Chanos) or a long 
tube-containing ossicle (Kneria) above it. The lower preopercular limb, however, 
runs well forward and forms the chief support for the quadrate. On the skull roof 
the frontals are large and the parietals are very small and almost restricted to the 
rear border of the skull, with the supraoccipital separating those of the two sides. 
The supratemporal commissure of the lateral line either runs across the parietals 
and supraoccipital through a series of superficial ossicles or through these bones 
themselves. 

In Gonorhynchus and Chanos the exoccipitals send out a pair of prongs that roof 
the anterior portion of the spinal cord ; in the flat-headed Kneria and Phractolaemus 
the projection of the supraoccipital serves the same purpose, with the walls of the 
first few neural arches forming the sides. 



354 A CLASSIFICATION OF MODERN ISOSPONDYLOUS FISHES 

The lower intermuscular bones are present in Chanos and Gonorhynchus , apparently 
absent in Phractolaemus and Kneria. In Gonorhynchus the lower ribs are short 
and slight, and articulate with lateral expansions of the centra. In Chanos the ribs 
are large and their upper portions articulate directly with the centra, the lower by 
way of a bony nodule. In Phractolaemus and Kneria the ribs articulate directly 
with the centra but in the former the ribs are very strong, in the latter moderate. 
Admittedly, these vertebral characters show little relationship between the genera, 
but they are given because of their previous use in classification. 

The caudal skeleton of all these groups {Cromeria}) is essentially the same. 

As to mode of life, Gonorhynchus is today a purely marine fish, but the group to 
which it belongs is known from Eocene fresh-water deposits from America and 
Europe. Chanos is euryhaline, and the other three genera are restricted to fresh 
water. All five genera are tropical. 

A good many of the similarities in these five groups may be primarily or secondarily 
related to the small size of the mouth in all these forms. It could therefore be that 
the small mouth has developed independently two or more times and that all of the 
other changes have followed as a matter of course. Some check on this possibility 
is offered by the members of the Argentinoidae, a small-mouthed group in the 
salmonoid stock. To some extent a parallel evolution is indicated between the 
Argentinoidae and the five groups taken up here, particularly with regard to jaw 
structure, suspensorium and the reduction in branchiostegal rays. However, there 
are a number of features in the five groups that are not duplicated in the Argentinoidae. 
Among these are the attachment of the gill covers to the isthmus, the loss of the 
normal upper portion of the preoperculum, the passage of the supratemporal commis- 
sure over or through the supraoccipital, the expansion of the anterior portion of 
the spinal canal, and the advanced form of the caudal skeleton. 

To summarize, the argument presented here is not that Gonorhynchus, Chanos, 
Cromeria, Kneria, and Phractolaemus are closely related. Indeed, no two of them 
are. It merely seems to the author that each of the five is more closely related to 
the others than it is to anything else. If this is granted, then it becomes fairly easy 
to tie in the group, via Chanos and to some extent Gonorhynchus, with the general 
clupeoid stock 1 . 

SECTION 4. A CLASSIFICATION OF THE MODERN 
ISOSPONDYLOUS FISHES 

The systematics of the living isospondylous fishes have received a great deal of 
attention. Because of the excellent literature review by Wilimovsky (1951) only a 
sampling of papers on the subject need be mentioned here. 

With regard to the general classification of the groups of Clupeiformes the deserved- 
ly classical papers are those of Ridewood (1904^ and b, 1905a, b, and c). Unfortu- 

1 The British Museum (Natural History) material upon which the above account has been based 
consists of complete skeletons of Gonorhynchus and Chanos, somewhat incomplete skeletons of Practolaemus 
and Kneria, and a stained and partially cleared skeleton of Kneria. Whole specimens of all genera 
mentioned have been examined. For the osteology of Cromeria, reliance has had to be placed on the 
paper by Swinnerton (1903). Ridewood's accounts of the head skeletons of Phractolaemus (1905a) and 
Gonorhynchus ( 19056) have also been used. 



A CLASSIFICATION OF MODERN ISOSPOND YLOUS FISHES 



355 



nately Ridewood studied only the head skeletons of a limited number of isospondy- 
lous groups. Regan (1913a) dealt briefly with the classification of the salmonoid 
fishes and in 1929 presented a synopsis of the full order. Berg's (1940) volume 
introduced some rather drastic innovations. Chapman (1934-1948) has written a 
number of valuable papers on the relationships of individual members of the group, 
and Norman (1957) gave a key to the suborders, families, and genera. 

Besides the above and many other papers dealing more or less directly with 
isospondylous classification, a large number of works bear indirectly on the subject. 
For example, Ford (1937) took up variations in the vertebral column. Whitehouse 
(1910a and b), Regan (1910a and b), Hollister (1936, 1940), and Schaeffer (1949 : 
13, 14) have taken up the caudal skeleton. Starks (1930) wrote the most complete 
account of the pectoral girdle. Frost (1925) took up isospondylous otoliths. Allis 
(1904), Wohlfahrt (1937), and Nybelin (1957) have dealt with the lateral line system 
of the head. Van Dobben (1935), Hofer (1945), Tchernavin (1953), Giinther and 




Diagram 3 
Deckert (1953, etc.) and Kirkhoff (1958) have taken up various aspects of the func- 
tional morphology of the head. As to soft anatomy, Bridge (1900), de Beaufort 
(1909), Svetovidov (1950, etc.), Jones and Marshall (1952), and O'Connell (1955) 
investigated the air bladder. Boas (1880) and Senior (1907, etc.) reported on the 
heart. Jacobshagen (1912) and Suyehiro (1942) have made comparative studies 
of the digestive system . Finally, Hyrtl (1856), Lagler and Kraatz (1945), Kapoor 
(1954), and Takahashi (1957) have dealt with the epibranchial organ. 

Despite all of the work that has been done, the classification of the isospondylous 
fishes (Order Clupeiformes) remains unsatisfactory. Indeed, Berg (1940 : 417) 
has written : " This order, from which a series of higher orders has arisen, represents 
an artificial assemblage, its separate members, as may be seen from the diagnosis, 
differing greatly from one another. In time the Clupeiformes will be, doubtlessly, 
divided in many orders ". Berg's view, I think, stems at least in part from a tendency 
to look at isospondylous classification at a single time level, for example AB of 
Diagram 3, whereas if a time dimension, CD, were incorporated into the concept, 
the " artifical assemblage " aspect would seem to largely disappear. In any event 
there are four main questions that need to be answered with regard to isospondylous 
classification. (1) What are the interrelationships of the living members? (2) What are 
the relationships between living and fossil members? (3) What are the limits of the 
group? (4) How can it be defined? 



356 A CLASSIFICATION OF MODERN ISOSPOND YLOUS FISHES 

Here, no effort will be made to deal with fossil forms. The classification of fossil 
Clupeiformes is a large and important subject in itself, and one to which the present 
author can make little contribution as he has no first hand familiarity with fossil 
material. 

The classification of modern isospondylous fishes that follows thus attempts to 
reconstruct phylogenetic relationships from a knowledge of fishes at only one time 
level and is consequently prone to all the pitfalls inherent in such a procedure. Yet 
there are certain points to be made in justification of such a classification. One is 
that it provides a corpus for the paleontologists to work from, with, or even against. 
More important, however, is the fact that ichthyologists working with modern 
fishes are obligated to do as much as they can with the fishes around them. There is 
virtually no chance that neotenic fishes like Cromeria and Lovettia will ever turn 
up in the fossil record. Indeed one is inclined to wonder whether the fossil record 
would not give a very peculiar picture of isospondylous classification because of 
the overrepresentation of some groups and the underrepresentation of others. 
Thus the salmonids are usually stream dwellers and are unlikely candidates for 
fossilization ; the same might be said of the oceanic stomiatoids. Again, some of 
the most peculiar of the isospondylous fishes are from African fresh waters, and 
pre-Pliocene fish remains from that continent are as yet few. On the other hand 
certain of the isospondylous fishes, notably the herring-like forms, seem frequently 
to die in areas where they fossilize in relative abundance. Thus if the major groups 
of the paleontologist, e.g., the chirocentrids, do not always correspond to the major 
groups of the modern ichthyologist, that may merely mean a difference in viewpoint 
rather than an error in interpretation. Eventually, of course, a single classification 
of isospondylous fishes based on both fossil and recent material will have to be 
developed, but the time to attempt this hardly seems to have arrived. 

As to a hard and fast definition of the Clupeiformes, that, as might be expected 
of any group so diverse, is impossible except in negative terms. Apparently about 
the best that can be done is as follows : 

Caudal fin homocercal. No true spines in the fins. Pelvic fins abdominal and with 
more than 5 rays (although in the Notopteridae the pelvics are absent or rudimentary 
and few-rayed, and the Notopteridae, Pantodon and Macristium have subthoracic 
pelvics). Maxillary usually included in the gape. Scales, if present, cycloid (except 
Gonorhynchus and certain species of argentinids and osmerids). Air bladder, if 
present, physostomous (except Lovettia, stomiatoids, and Argentinoidae) . Orbitos- 
phenoid and mesocoracoid generally present ; branched caudal rays most 
frequently 17. No Weberian ossicles. Lateral line, if present, not running low on 
sides. Snout not tubular. No electric organs. 

With regard to the limits of the Clupeiformes, both the " upper " and the " lower " 
borders of the order have been the subject of controversy. Thus Saint-Seine (1949) 
removed Elops from the isospondylous fishes and placed it in the Halecostomi, 
and Nybelin (1957) has stated that Elops is as much a holostean as Amia or Lepiso- 
steus. Here, Elops will be retained in the Clupeiformes, with the full realization that 
it becomes extremely difficult to place a lower limit on the order when fossil material 
is taken into account. 



A CLASSIFICATION OF MODERN ISOSPOND YLOUS FISHES 357 

At the " upper " limit there are six groups of fishes that have been at times included 
in and at other times excluded from the Clupeiformes by modern authors. That 
the Anotopteridae, included " incertae sedis " by Berg (1940: 435), belongs 
among the iniomous fishes has been adequately shown by Maul (1946) and Nybelin 
(1946). Berg (1940 : 436) creates a separate order for the Galaxiidae, but this family 
may apparently be returned to its usual place in the Clupeiformes (see Section 2). 
The Bathyclupeidae has been juggled about by various authors, sometimes landing 
in the isospondylous fishes, but it belongs with the percoids where Regan (1913&) 
placed it (N. B. Marshall). The three remaining groups are more controversial. 
The gymnarchid-mormyrid stock can be traced back with considerable certainty 
into the osteoglossid-notopterid group of isospondylous fishes. Nevertheless the 
Mormyriformes seems to show sufficient specializations — e.g., in the electric organ 
and the brain — to justify separation at the ordinal level. The controversy regarding 
the iniomous fishes has a somewhat different basis. Their origin in the Clupeiformes 
cannot be traced by more than guesswork. However, the one character usually 
used to differentiate the iniomous from the isospondylous fishes is the exclusion of 
the maxillary from the gape in the former, and Chapman (1944) and others have 
pointed out that various isospondylous families also have the maxillary excluded. 
Thus the differentiation between the two orders breaks down, and Schultz and Stern 
(1948) have consequently merged them. Nevertheless, it does not seem justifiable 
to unite two groups just because all themembers of one cannot be separated from 
all the members of the other by a single character. In the instance of the Mycto- 
phiformes and the Clupeiformes it is not even a question of where to draw the line 
in a continuous lineage, for the isospondylous fishes with the maxillary excluded 
from the gape (except possibly Albula) undoubtedly did not give rise to the iniomous 
fishes. But if the argument for merging the two orders seems weak, that for keeping 
them separate as is done here has, in this author's opinion, never been very convinc- 
ingly presented. (The closest approach seems to be that of Marshall, 1955 : 305). 
Finally, that the haplomous fishes belong in the Clupeiformes has been convincingly 
argued by Berg (1936, 1940), and further evidence for this allocation has already 
been presented in Section 2 of this paper. 

Order Clupeiformes (Isospondyli) 
Division I Clupei 

Suborder I Elopoidei 

Superfamily I Elopoidae 

Superfamily II Albuloidae 
Suborder II Clupeoidei 

Superfamily I Alepocephaloidae 

Superfamily II Clupeoidae 
Suborder III Gonorhynchoidei 

Superfamily I Chanoidae 

Superfamily II Gonorhynchoidae 

Superfamily III Phractolaemoidae 

Superfamily IV Knerioidae 

Superfamily V Cromerioidae 



358 A CLASSIFICATION OF MODERN ISOSPOND YLOUS FISHES 

Suborder IV Stomiatoidei 

Superfamily I Gonostomoidae 

Superfamily II Stomiatoidae 

Superfamily III Astronesthoidae 
Suborder V Salmonoidei 

Superfamily I Salmonoidae 

Superfamily II Argentinoidae (Opisthoproctoidae) 

Superfamily III Osmeroidae 
Suborder VI Esocoidei (Haplomi) 

Superfamily I Dallioidae 

Superfamily II Umbroidae 

Superfamily III Esocoidae 
Division II Osteoglossi 

Suborder VII Osteoglossoidei 

Superfamily I Hiodontoidae 

Superfamily II Notopteroidae 

Superfamily III Osteoglossoidae 

Certain final points regarding the classification of the isospondylous fishes pre- 
sented above may be noted. First, N. B. Marshall has called my attention to the 
omission of three families of dubious systematic position : the Bathylaconidae 
(Parr, 1948), the Rosauridae (Tucker, 1954) and the Macristiidae (Regan, 1911). 
These are all families of oceanic fishes based on one or a few specimens. Any attempt 
to place them in one position or another within the Clupeiformes at the present time 
would be gratuitous, and it seems better to leave them incertae sedis. As to the better- 
represented groups, the classification of the superfamilies of Stomiatoidei and Eso- 
coidei stand as they appear in Berg (1940), and the superfamily Alepocephaloidae is 
left as usual in the Clupeoidei. The author has almost no first hand knowledge of 
any of these groups. 

Second, as a matter of practice this author believes that orders and suborders 
should be monophyletic but rather broadly interpreted. In contrast with Berg 
(1940), he would be very chary about the erection of orders and suborders for a 
few aberrant forms ; these he would in general relegate to superfamilies (if such a 
higher category is needed) . The reasoning behind this is that superfamilies are usually 
used by specialists, to whom the names of small divergent groups are significant, 
but are rarely incorporated in general classifications, where such names would 
generally have little meaning. 

Finally, there remains the question of whether the Clupeiformes is polyphyletic. 
The evidence of the caudal skeleton would appear to be against such a conclusion. 
The caudal structures of the basal forms Elops, Salmo, Esox, and Hiodon are not only 
strikingly similar (Section 1), they are also widely different from anything among 
the recent members of the Holostei. Some of the ways in which they resemble 
each other and differ from the caudal skeletons of lower fishes seem to be of a type that 
would hardly have been developed independently twice, e.g., the three and only 
three hypurals to the lower caudal lobe, the peculiarities in the basal articulations 



A CLASSIFICATION OF MODERN ISOSPOND YLOUS FISHES 359 

of these three hypurals (No. 1 with the terminal vertebra, and Nos. 2 and 3 with the 
first postterminal centrum), the long uroneurals, etc. Indeed if there is any indication 
of polyphyletic origin in the caudal structure of the Clupeiformes it would be between 
the osteoglossid-notopterid stock on the one hand and Hiodon with all the rest of 
the isospondylous fishes on the other. But such a split separating Hiodon from 
the osteoglossid groups seems untenable on any other ground than the caudal skeleton. 

When all features are taken into consideration only two stocks (Divisions) seem 
distinctly separable within the order. One of these (Division II) contains today 
a small number of relict forms that are all, in their peculiar fashion, mixtures of 
primitive and specialized characters. To exemplify the basic nature of the stock 
no one fish can be selected ; rather it is necessary to give a composite description 
based on the primitive features found in most of the Hiodontoidae, Osteoglossoidae, 
and Notopteroidae. If this is done it may be said that in Division II the parasphenoid 
remains primitive from three points of view. First, it never reaches the rear of the 
basioccipital ; second, it sometimes retains a pair of projections for articulation 
with the suspensorium ; and finally, it generally bears teeth. The last two features 
play a role in the general dentition and palatal construction in the stock as a whole ; 
in these Division II appears to have remained primitive, quite in contrast to the 
evolution that has occurred in Division I. Throughout Division II, except Heterotis, 
the primary portion of the bite is between the parasphenoid and the tongue (cf., 
Hofer, 1945). Such bony articulations as occur on the top of the mouth are between 
the parasphenoid and the mesopterygoid, not from the ethmoid-prefrontal area 
via the palatines to the maxillary pedicels. Indeed, in Division II the palatines 
end anteriorly in a simple point and the maxillaries merely appear to be toothed 
cheek bones that have come loose posteriorly ; they have no pedicels. However, 
if supramaxillaries are a primitive character, Division II has lost them. Also, in 
the vertebral column Division II would seem to have advanced farther along the 
road toward the fusion of elements than Division I, and in the caudal skeleton 
(aside from Hiodon) Division II seems to have evolved in a totally different way 
from Division I. 

By contrast Division I is represented today by some primitive forms, notably 
the elopoids, that in most respects might stand as the ancestors of the whole division. 
Actually, Elops and Albula in certain characteristics, especially dentition, seem to 
form better prototypes for the stock than the Jurassic Leptolepis bronni (Rayner, 
1937), for in dentition L. bronni seems already to have developed much of the speci- 
alization of the modern Clupeidae. Returning to modern forms, the Elopoidae and 
Albuloidae are the only superfamilies in the Division that have retained a para- 
sphenoid dentition. In the others it has been totally lost, and when teeth reappear 
along the center of the mouth roof as in Esox they are on a backward projection of 
the vomer. The parasphenoid-mesopterygoid articulation is also lost in all modern 
forms though it is present in Leptolepis bronni (Rayner, 1937). On the other hand, 
a complicated cranium-palatine-maxillary articulation is usually present, giving 
the whole upper jaw structure an aspect rather similar to that of the higher fishes. 

An important question regarding Division I is whether it contains within itself 
two or more independent lineages that are of equal value with Division II. One 



360 A CLASSIFICATION OF MODERN ISOSPOND YLOUS FISHES 

such possible independent group is the Gonorhynchidae. If, however, Gonorhynchus 
is related to Chanos, as is suggested in Section 3 of this paper, then its pertinence 
to Division I would seem fairly secure. A more moot group, in this author's opinion, 
is the whole salmonoid-haplomous stock. There are only two items known to the 
author that might serve as an indication of a salmonoid-clupeoid relationship. 
One is that Albula, which is usually assumed to be related to the clupeoids, seems 
to show some similarity in the caudal skeleton to that typical of the salmonoids. 
The other is that the stomiatoids appear to be intermediate in certain respects 
between the salmonoids and the clupeoids : on the one hand some of the stomiatoids 
have the two supramaxillaries of the clupeoid fishes and Elops ; on the other, many 
stomiatoids have the typical salmonoid adipose fin. Especially because of this 
puzzling link provided by the stomiatoids, it seems best to include the salmonoids 
in the same division with the clupeoids for the moment. 



Synopsis of Modern Clupeiform Fishes to Superfamily 

1 a. Parasphenoid extending nearly to the posterior end of the basioccipital or beyond. 

Parasphenoid never with lateral processes for articulation with the mesoptery- 

goid ; anterior ribs usually articulating with small parapophyses that are wedged 

into pits in the centra ; branched caudal rays often 17. Division Clupei. 
2a. Teeth present on the parasphenoid ; a leptocephalous larval form. 

Suborder Elopoidei 
3a. A gular plate ........ Superfamily Elopoidae 

3b. No gular plate ........ Superfamily Albuloidae 

2b. No teeth on the parasphenoid ; no leptocephalous larval form. 

4a. Lower intermuscular bones usually present ; photophores, if present, not in two rows 

along lower sides ; posterior neural and hemal spines without antero-posteriorly 

expanded blades ; no proethmoids. 
5a. Supramaxillaries usually present ; gill covers free from the isthmus ; branchio- 

stegal rays 7 or more ....... Suborder Clupeoidei 

6a. Two postterminal centra ; no temporal foramen ; black, deep-water fishes 

Superfamily Alepocephaloidae 
6b. One or no postterminal centra ; a temporal foramen ; silvery, surface-living 

fishes ......... Superfamily Clupeoidae 

5b. No supramaxillaries ; gill covers attached to the isthmus ; branchiostegal rays 

3 or 4 . . . . . . . Suborder Gonorhynchoidei 

7a. Scales cycloid ; gill openings not restricted ; anterior ribs articulating in part with 

parapophyses wedged into pits in the centra ; mouth terminal 

Superfamily Chanoidae 
7b. Scales ctenoid ; gill openings not restricted ; anterior ribs articulating with strong 

lateral processes from the centra ; mouth inferior 

Superfamily Gonorhynchoidae 
7c Scales cycloid ; gill openings little restricted ; anterior ribs articulating directly 

with the centra ; mouth superior . . . Superfamily Phractolaemoidae 

7d. Scales cycloid ; gill openings restricted ; anterior ribs articulating directly with 

the centra ; mouth inferior ..... Superfamily Knerioidae 

7c No scales ; gill openings restricted ; mouth inferior . Superfamily Cromerioidae 

4b. No lower intermuscular bones ; two rows of photophores along lower sides ; 

posterior neural and hemal spines without antero-posteriorly expanded blades ; 

no proethmoids ....... Suborder Stomiatoidei 



A CLASSIFICATION OF MODERN ISOSPOND YLOUS FISHES 361 

8a. Premaxillary without anterior expansion ; maxillary with two supramaxillaries ; 

gill arches with gill rakers ..... Superfamily Gonostomoidae 

8b. Premaxillary with a strong anterior expansion extending upward over the ethmoid; 

maxillary without supramaxillaries ; gill arches without normal gill rakers. 
9a. Supracleithrum absent ....... Superfamily Stomiatoidae 

9b. Supracleithrum present ..... Superfamily Astronesthoidae 

4c. No lower intermuscular bones ; no photophores ; posterior neural and hemal spines 
with antero-posteriorly expanded blades (except in Lovettia and the Salangidae) ; 
proethmoids sometimes present .... Suborder Salmonoidei 

10a. Two upturned postterminal vertebral centra . . . Superfamily Salmonoidae 

10b. Not more than one upturned postterminal vertebral centrum. 

11a. Oceanic fishes with small pelagic eggs. . . . Superfamily Argentinoidae. 

11b. Inshore or freshwater fishes with large, demersal eggs . Superfamily Osmeroidae 

4d. No lower intermuscular bones, photophores, or antero-posteriorly expanded blades 
on the posterior neural and hemal spines; paired proethmoids present 

Suborder Esocoidei 
12a. Snout not greatly produced ; no canine teeth. 

13a. Scapula, coracoid, and pectoral radials not ossified . . Superfamily Dallioidae 

13b. Scapula, coracoid, and pectoral radials ossified . . Superfamily Umbroidae 

12b. Snout greatly produced ; canine teeth present. . . Superfamily Esocoidae 

ib. Parasphenoid terminating well short of the posterior end of the basioccipital. 
Parasphenoid sometimes with lateral processes for articulation with the meso- 
pterygoid ; anterior ribs either articulating directly with centra or with strong 
parapophyses that are fused to the centra ; branched caudal rays 16 or fewer 

Division Osteoglossi 
14a. Three or four uroneurals in the caudal skeleton ; no parasphenoid facet for 
articulation with the mesopterygoid ; nasals small ; symplectic a normal, splint- 
like bone ; basisphenoid absent ; top of skull without longitudinal ridges. 

Superfamily Hiodontoidae. 
14b. No separate uroneurals nor parasphenoid facet ; nasals enlarged ; symplectic 
expanded ; basisphenoid present ; top of skull with longitudinal ridges 

Superfamily Notopteroidae 
14c. No separate uroneurals ; parasphenoid facets present ; nasals enlarged ; basi- 
sphenoid present ; top of skull rugose but without longitudinal ridges 

Superfamily Osteoglossoidae 

REFERENCES 

Numbers in parentheses after each item refer to the section of the present paper in which 
this item was cited. 

Allis, E. P. 1904. The latero-sensory canals and related bones in fishes. Intern. Monats. 

Anat. Physiol., 4 : 401-502, pis. 8-20. (4) 
Barrington, E. J. W. 1937. The structure and development of the tail in the plaice (Pleuro- 

nectes platessa) and the cod (Gadus morrhua). Quart. Jour. Micr. Sci., London, new series, 

79 : 447-469, 25 figs. (1) 
de Beaufort, L. F. 1909. Die Schwimmblase der Malacopterygier. Morph. Jahrb., 39 : 529- 

644, 1 pi., 10 figs. (4) 
Berg, L. S. 1936. The suborder Esocoidei (Pisces). Bull. Inst. rech. biol. Perm, 10 : 385-391. 

(In Russian and English). (2, 4) 
1940. Classification of fishes, both recent and fossil. Trav. Inst. zool. Acad. Sci. U.R.S.S., 

Leningrad, 5 : 87-517, 190 figs. (In Russian and English). (2, 3, 4) 

1955- The same. 2nd Ed. Ibid., 20 : 1-286, 263 figs. (In Russian). (2) 

Bertelsen, E. 1958. The argentinoid fish Xenophthalmichthys danae Regan 1925. Dana- 
Report No. 45, pp. 3-8, 6 figs. (2) 



362 A CLASSIFICATION OF MODERN ISOSPOND YLOUS FISHES 

Blackburn, M. 1950. The Tasmanian whitebait, Lovettia seali (Johnston), and the whitebait 

fishery. Australian Jour. Mar. and Freshw. Res., 1 : 155-198, 6 pis., 5 figs. (2) 
Blanco, G. J. 1935. The development of the homocercal caudal of the blue perch, Taeniotoca 

lateralis Agassiz. Philippine Jour. Sci., Manila, 56 : 41-44, 1 pi. (1) 
Boas, J. E. V. 1880. Uber den Conus arteriosus bei Butirinus und anderen Knochenfischen. 

Morph. Jahrb., 6 : 527-531, 1 pi. (4) 
Bridge, T. W. 1900. The air-bladder and its connection with the auditory organ in Notop- 

terus borneensis. Jour. Linnaean Soc. London, Zool., 27 : 503-540, 2 pis. (4) 
Chapman, W. M. 1934. The osteology of the haplomous fish, Novumbra hubbsi Schultz, 

with comparative notes on related species. Jour. Morph., 56 : 371-405, 8 figs. (2, 4) 

1941. The osteology and relationships of the isospondylous fish Plecoglossus altivelis 

Temminck and Schlegel. Ibid., 68 : 425-455, 12 figs. (4) 

1942a. The osteology and relationships of the Argentinidae, a family of oceanic fishes. 

Jour. Washington Acad. Sci., 34 : 104-117, 8 figs. (2, 4) 

1 942ft. The osteology and relationships of the bathypelagic fish Macropinna microstoma, 

with notes on its visceral anatomy. Ann. Mag. Nat. Hist., ser. 11, 9 : 272-304, 9 figs. (1, 

2,4) 

1943- The osteology and relationships of the bathypelagic fishes of the genus Bathylagus 

Giinther with notes on the systematic position of Leuroglossus stilbius Gilbert and Thero- 
bromus callorhinus Lucas. Jour. Washington Acad. Sci., 33 : 147-160, 8 figs. (2, 4) 

1944- On the osteology and relationships of the South American fish, Aplochiton zebra 

Jenyns. Jour. Morph., 75 : 149-165, 10 figs. (1, 2, 4) 

1948. The osteology and relationships of the Microstomidae, a family of oceanic fishes. 

Proc. California Acad. Sci., ser. 4, 26 : 1-22, 12 figs. (1, 2, 4) 

Cohen, D. M. 1958. A revision of the fishes of the subfamily Argentininae. Bull. Florida 

State Mus., 3 : 93-173, 8 figs. (2) 
Devillers, C. 1958. Le crane des poissons. In Grass e, Traite de Zoologie, vol. 13, fasc 1, 

pp. 551-587. ngs. 345-443- Paris. (1) 
Eigenmann, C. H. 1927. The freshwater fishes of Chile. Mem. Nat. Acad. Sci., 22 : no. 2, 

63 pp., 16 pis. (2) 
Emelianov, S. 1935. Die Morphologie der Fischrippen. Zool. Jahrb., Abt. Anat., 60 : 133-262, 

53 figs- (2) 
Ford, E. 1937. Vertebral variation in teleostean fishes. Jour. Mar. Biol. Ass., new ser., 22 : 

1-60, pis. 1-16, 18 figs. (4) 
Frost, G. A. 1925. A comparative study of the otoliths of neopterygian fishes. Ann. Mag. 

Nat. Hist., ser. 9, 15 : 152-163, pis. 11-13. (4) 
Garstang, W. 1931. The phyletic classification of Teleostei. Proc. Leeds Philos. Lt. Soc, 

Sci. Sect., 2 : pt. 5, pp. 240-260. (2) 
Goodrich, E. S. 1909. Cyclostomes and Fishes. Vol. 9 of Lankester's " Treatise on Zoology ". 

Adam and Charles Black, London : xvi + 518 pp., 515 figs. (1) 
Greene, C. W. & Greene, C. H. 1914. The skeletal musculature of the king salmon. Bull. 

U.S. Bur. Fish, for 1913, 33 : 21-60, pis. 1, 2, 14 figs. (1) 
GtJNTHER, K. & Deckert, K. 1953- Morphologisch-anatomische und vergleichend okolo- 

gische Untersuchungen fiber die Leistungen des Viszeralapparates bei Tiefseefischen der 

Gattung Cyclothone (Teleostei, Isospondyli) . Zeitschr. Morph. Okol., 42 : 1-66, 25 figs. (4) 
Hindersson, H. A. 1910. Tiber die Schwanzflossenmuskulatur der Teleostier. Anat. Anz., 

36 : 465-471. 5 figs- (1) 
Hofer, H. 1945. Zur Kenntnis der Suspensionsformen des Kieferbogens und deren Zusam- 

menhange mit dem Bau des knochernen Gaumens und mit der Kinetik des Schadels bei 

den Knochenfischen. Zool. Jahrb., Abt. Anat., 69 : 321-404, 21 figs. (4) 
Hollister, G. 1936. Caudal skeleton of Bermuda shallow water fishes. I. Order Isospondyli : 

Elopidae, Megalopidae, Albulidae, Clupeidae, Dussumieriidae, Engraulidae. Zoologica, 

New York, 21 : 257-290, 53 figs. (1, 4) 



A CLASSIFICATION OF MODERN ISOSPOND YLOUS FISHES 363 

Hollister, R. G. 1940. Young Megalops cyprinoides from Batavia, Dutch East India, includ- 
ing a study of the caudal skeleton and a comparison with the Atlantic species, Tarpon 
atlanticus. Ibid., 24 : 449-467, 21 figs. (1,4) 

Hubbs, C. L. 1953. Synonymy of the bathypelagic fish genus Rhynchohyalus, referred to 
the expanded family Argentinidae. Copeia, 1953, pp. 96-97. (2) 

Hyrtl, C. J. 1856. Uber die accessorischen Kiemenorgane und den Darmcanal der Clupea- 
ceen. Denkschr. Akad. Wiss. Wien, math.-nat. CI., 10 : 47-58, 3 pis. (4) 

Jacobshagen, E. 191 2. Untersuchungen iiber das Darmsystem der Fische und Dipnoer. 
Teil II. Materialien zur vergleichenden Anatomie des Darmkanals der Teleostomen nebst 
einer einleitenden U bersicht. Jenaische Zeitschr. Naturwiss., 49 : 373-810, pi. 18, 164 
figs. (4) 

Jones, F. R., & Marshall, N. B. 1953. The structure and functions of the teleostean swim- 
bladder. Biol. Rev., 28 : 16-83, 7 figs. (4) 

Kapoor, B. G. 1954. The pharyngeal organ and its associated structures in the milk-fish, 
Chanos chanos (Forskal). Jour. Zool. Soc. India, 6 : 51-58, 5 figs. (4) 

Kendall, W. C. 192 i. Peritoneal membranes, ovaries, and oviducts of salmonoid fishes 
and their significance in fish-cultural practices. Bull. Bur. Fish., Washington, 37 : 185-208, 
11 figs. (2) 

1935- The fishes of New England. The Salmon family. Part 2 — The Salmons. Mem. 

Boston. Soc. Nat. Hist., 9 : 1-166, 11 col. pis. (2) 

Kendall, W. C. & Crawford, D. R. 1922. Notice of a spiral valve in the teleostean fish 

Argentina silus, with a discussion of some skeletal and other characters. Jour. Washington 

Acad. Sci., 12 : 8-19, 2 figs. (2) 
Kirkhoff, H. 1958. Functionell-anatomische Untersuchung des Visceralapparates von 

Clupea harengus L. Zool. Jahrb., Abt. Anat., 76 : 461-570, 63 figs. 
Kolliker, A. i860. Uber das Ende der Wirbelsaule der Ganoiden und einiger Teleostier. 

Leipzig : 27 pp., 4 pis. (1) 
Lagler, K. F. & Kraatz, W. C. 1945. Pharyngeal pockets in the gizzard shad, Dorosoma 

cepedianum (Lesueur). Pap. Michigan Acad. Sci., Arts, Lett., 30 : 311-320, 2 pis., 1 fig. (4). 
Lotz, T. 1864. Uber den Bau der Schwanzevirbelsaule der Salmoniden, Cyprinoiden, Per- 

coiden und Cataphracten. Zeitschr. Wiss. Zool., 14 : 81-106, 4 pis., 2 figs. (1) 
Marshall, N. B. 1955. Alepisauroid fishes. Discovery Repts., 27 : 303-336, pi. 19, 9 figs. (4) 
Maul, G. E. 1946. Monografia dos peixes do Museu Municipal do Funchal. Ordem Iniomi. 

Bol. Mus. Mun. Funchal, No. 2, art. 2, pp. 5-61, 22 figs. (4) 
Norman, J. R. 1957. A draft synopsis of the orders, families and genera of recent fishes and 

fish-like vertebrates. British Museum (Natural History), London : 649 pp. (4) 
Nybelin, O. 1946. Sur quelques especes du genre Anotopterus Zugmayer et sur la position 

systematique de la famille des Anotopteridae. Goteborgs Vetensk. Samh. Handl., ser. 6, 

5b : 3-16, 1 pi. (4) 

1957- Les canaux sensoriels du museau chez Flops sauriis (L.) Notice preliminaire. 

Arkiv for Zoologi, 10 : 453-45$, 3 figs- (4) 

O'Connell, C. P. 1955. The gas bladder and its relation to the inner ear in Sardinops caerulea 
and Engraulis mordax. Bull. U.S. Fish Wildlife Serv., 56 : 503-553, 15 figs. (4) 

Parr, A. E. 1937. Concluding report on fishes. Bull. Bingham Oceanogr. Coll., 3, art. 7, 79 pp., 
22 figs. (2) 

1948. The classification of the fishes of the genera Bathylaco and Macromastax, possible 

intermediates between the Isospondyli and the Iniomi. Copeia, 1948, pp. 48-54, 2 figs. (4) 

Ramanujam, S. G. M. 1929, The study of the development of the vertebral column in teleosts, 

as shown by the life history of the herring. Proc. Zool. Soc. London, 1929, pp. 365-414, 

28 figs. (1) 
Rayner, D. N. 1937. On Leptolepis bronni Agassiz. Ann. Mag. Nat. Hist., ser. 10, 19 : 

46-74, 14 figs. (4). 
Regan, C. T. 1905. A revision of the fishes of the family Galaxiidae. Proc. Zool. Soc. London 

1905, pp. 363-384* 4 Pis. (2) 



364 A CLASSIFICATION OF MODERN ISOSPOND YLOUS FISHES 

Regan, C. T. 1910a. The caudal fin of the Elopidae and of some other teleostean fishes. Ann. 
Mag. Nat. Hist., ser. 8, 5 : 354-358, 2 figs. (1,4) 

19106. On the caudal fin of the Clupeidae, and on the teleostean urostyle. Ibid., pp. 

531-533. 2 figs. (1, 4) 

191 1. On the systematic position of Macristium chavesi. Ibid., 7 : 204-205, 1 fig. (4) 

191 3fl. The antarctic fishes of the Scottish National Antarctic Expedition. Trans. Roy. 

Soc. Edinburgh, 49 : 229-291, 11 pis., 6 figs. (2, 4) 

19136. The classification of the percoid fishes. Ann. Mag. Nat. Hist., ser. 12, 8 : 111-145. (4) 

■ 1929. " Fishes " in Encyclopaedia Britannica, ed. 14, 9 : 305-328. (2) 

Ridewood, W. G. 1904a. On the cranial osteology of the fishes of the families Elopidae and 

Albulidae, with remarks on the morphology of the skull in the lower teleostean fishes 

generally. Proc. Zool. Soc. London, 1904, pt. 2, pp. 35-81, figs. 8-18. (4) 
1904^. On the cranial osteology of the fishes of the families Mormyridae, Notopteridae 

and Hyodontidae. Jour. Linnaean Soc. London, Zool., 29 : 188-217, pis. 22-25. (4) 
1905a. On the cranial osteology of the clupeoid fishes. Proc. Zool. Soc. London, 1904, 

pt. 2, pp. 448-493, figs. 1 18-143. (4) 
19056. On the cranial osteology of the fishes of the families Osteoglossidae, Pantodontidae 

and Phractolaemidae. Jour. Linnaean Soc. London, Zool., 29 : 252-282, pis. 30-32. (4) 

1905c. On the skull of Gonorhynchus greyi. Ann. Mag. Nat. Hist., ser. 7, 15 : 361-372, 

pi. 16, 1 fig. (3, 4) 

Saint-Seine, P. de. 1949. Les poissons des calcaires lithographiques de Cerin (Ain). Nouv. 

Arch. Mus. Hist. nat. Lyon, Fasc. 2, pp. 1-357, 2D pl s -> I2 ° n g s - (4) 
Schaeffer, B. 1 949. A teleost from the Livingston formation of Montana. American 

Mus. Novitates No. 1427, 15 pp., 2 figs. (4) 
Schmalhausen, I. I. 191 3. Zur Morphologie der unpaaren Flossen. II. Bau und Phylogenese 

der unpaaren Flossen und insbesonders der Schwanzrlosse der Fische. Zeitschr. Wiss. 

Zool., 104 : i-8o, 14 figs. 
Schultz, L. P. & Stern, E. M. 1948. The ways of fishes. I). Van Nostrand, New York : 

264 pp. (2, 4) 
Scott, E. O. G. 1936. Observations on fishes of the family Galaxiidae. Pt. 1. Pap. Proc. 

Roy. Soc. Tasmania, 1935, pp. 85-112, 4 figs. (2) 
Senior, H. D. 1907. Note on the conus arteriosus of Megalops cyprinoides (Broussonet). 

Biol. Bull, 12 : 378-379, 2 figs. (4) 
Starks, E. C. 1930. The primary shoulder girdle of the bony fishes. Stanford Univ. Ser., 

Biol. Sci., 6 : 149-239, 38 figs. (4) 
Stokell, G. 1941. A revision of the genus Retropinna. Rcc. Canterbury Mus., 7 : 361-372, 

Pis. 55-57- (2) 

1945- The systematic arrangement of the New Zealand Galaxiidae. Part I. Generic 

and sub-generic classification. Trans. Roy. Soc. New Zealand, 77 : 124-137, pis. 8-12. (2) 

1955- Fresh water fishes of New Zealand. Simpson and Williams, Christchurch : 145 pp., 

42 pis. (2) 
Suyehiro, Y. 1942. A study of the digestive system and feeding habits of fish. Japan. 

Jour. Zool., 10 : 1-303, 15 pis., 190 figs. (4) 
Svetovidov, A. N. 1950. Peculiarities in the structure of the air-bladder in Clupeidae. 

C. R. Acad. Sci. Moscou, n. s., 74 : 643-644. (In Russian). (4) 
Swinnerton, H. H. 1903. The osteology of Cromeria nilotica and Galaxias attenuatus. 

Zool. Jahrb., Abt. Anat., 18 : 58-70, 16 figs. (2) 
Takahashi, N. 1957. On the so-called accessory respiratory organ " gill helix " found in 

some clupeiform fishes, with special reference to the function and genealogy. Japan. Jour. 

Ichthy., 5 : 71-77, 1 pi. (4) 
Tchernavin, V. V. 1953. The feeding mechanisms of a deep sea fish Chauliodus sloani 

Schneider. British Museum (Natural History), London : 99 pp., 10 pis., 36 figs. (4) 
Totton, A. K. 1914. The structure and development of the caudal skeleton of the teleostean 

fish, Pleuragramma antarcticum. Proc. Zool. Soc. London, 1914, pp. 251-262, 2 pis. (1) 



A CLASSIFICATION OF MODERN I SOSPO X D YLOUS FISHES 365 

Trewavas, E. 1933. On the structure of two oceanic fishes, Cyema atrum Giinther and 
Opisthoproctas soleatus Vaillant. Proc. Zool. Soc. London, 1933, pp. 601-614, 2 V^ s -> 8 n g s - 

(1.2) 

Tucker, D. W. 1954. Tne " Rosaura " Expedition. 4. Report on the fishes . . . Part I. 

Bull. British Mus. {Nat. Hist.), Zool., 2 : 163-214, pis. 7, 8, 19 figs. (4) 
van Dobben, W. H. 1935- tJber den Kiefermechanismus der Knochenhschc. Arch. Nierland. 

Zool., 2 : 1-72, 50 figs. (4) 
Voigt, E. 1934. Die Fische aus der mitteleo/.anen Braunkohle des Geiseltales. Nova Acta 

Leopoldina, n. F., 2 : 22-146, 14 pis., 23 figs. (2) 
Whitehouse, R. H. 1910a. The caudal fin of the Teleostomi. Proc. Zool. Soc. London, 

1910, pp. 590-625, pis. 47-50, fig. 57. (1, 4) 
19106. Some remarks on the teleostean caudal fin. Ann. Mag. Nat. Hist., ser. 8, 5 : 425- 

427- (I. 4) 

1 91 8. The evolution of the caudal fins of fishes. Rec. Indian Mus., 15 : 135-142. (1) 

Wilimovsky, N. J. 1951. The classification of isospondylous fishes. A. S. I. H. Committee 

on Fish Classification, Circular No. 4, 38 pp. (Mimeographed). (2, 3, 4) 
Wohlfahrt, T. A. 1937. Anatomische Untersuchungen fiber die Seitenkanale der Sardine 

(Clupea pilchardus Walb.). Zeitschr. Morph. Okol. Tiere, 33 : 381-411, 15 figs. (4) 




THE ECOLOGY AND 

TAXONOMY OF SOME 

ANGOLA BIRDS 

(Based on a collection made in 1957) 




B. P. HALL 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ZOOLOGY Vol. 6 No. 7 

LONDON: i960 



THE ECOLOGY AND TAXONOMY OF SOME 

ANGOLA BIRDS 

(Based on a collection made in 1957) 



BY 

B. P. HALL 



Pp. 367-453 ; Plates 5-6 ; 2 Text-figures 




BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ZOOLOGY Vol. 6 No. 7 

LONDON: i960 



THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted in 1949, is 
issued in five series, corresponding to the Departments 
of the Museum, and an Historical series. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

This paper is Vol. 6, No. 7 of the Zoological series. 



© Trustees of the British Museum, i960 



PRINTED BY ORDER OF THE TRUSTEES OF 
THE BRITISH MUSEUM 

Issued July, i960 Price Forty Shillings 



THE ECOLOGY AND TAXONOMY OF SOME 

ANGOLA BIRDS 

(Based on a collection made in 1957) 

By B. P. HALL 

CONTENTS 

Page 
Introduction ........... 370 

Scope of paper 

Acknowledgments 

Deposition of specimens collected 
Itinerary ............ 372 

Vegetation ........... 373 

Main vegetational regions, with map 

Vegetation at each Camp 
Tables of Specimens Collected ....... 379 

Table I. Birds of the woodlands 
II. Birds of the forests 

III. Birds of grassland, reeds and open country .... 

IV. Birds of the thornbush semi-desert, tangled thickets and rocks 

of western districts 
Distribution Throughout Africa of Species found in Angola . . 390 

Woodland birds 

Forest birds 

Grassland birds 

Birds of the western districts 
Avifaunal Zones and Speciation ....... 395 

Avifaunal zones, with map 

Speciation 

Examples of stages in speciation 
Breeding Seasons .......... 401 

General discussion 

Some relevant rainfall figures 

Non-passerines 

Woodland passerines 

Forest passerines 

Grassland passerines 

Passerines of western districts 
New Records for Angola ......... 405 

Species not hitherto recorded in the country 

Extension of ranges 

New races which have been described on birds from the collection 

Sight records of interes 
Systematic and Field Notes on Selected Species .... 406 

Scope of the notes 

Notes on 84 species 
Bibliography ........... 450 

Appendix ............ 451 

List of Angola place-names mentioned in the text, other than camps 
zool. 6, 7. 22 



37o THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 

INTRODUCTION 

Since Bocage published the Ornithologie d' Angola in 1881 no major work has 
appeared on the birds of the country as a whole and even the vast collection amassed 
by Ansorge in the early part of the century has never been fully written up. Thus 
little knowledge has been available to the student on the ranges and habitats of 
the many forms endemic to Angola although the discoveries of such collectors as 
Rudyerd Boulton, Gerd Heinrich and Rudolf Braun in the past twenty-five years 
have shown it to be a country of exceptional interest. This interest lies not only 
in its endemic species but also in that it is a meeting place for the avifauna of West 
Africa, found in the lowland forests of the west ; of South West Africa, found in 
the coastal thorn country and desert ; of East Africa, in the woodlands of the 
centre and east. There are also some representatives of a specialised montane 
avifauna in the Bailundo highlands. 

Chapin's Birds of the Belgian Congo has done much to remedy the lack of knowledge 
but, of necessity, deals only briefly with Angola birds, and only with those that 
are found also in the Congo. In the British Museum workers have been handicapped 
by the paucity of specimens from Angola as well as by the lack of literature on the 
country, and it was with a view to filling some of the gaps in the National Collection 
and with learning something of the habitats and ecology of the birds, that I decided 
to make a collecting trip to Angola in 1957. 

This paper is intended to correlate the birds that were collected with the type of 
country and vegetation in which they were found. For this purpose all specimens 
have been listed in four "Tables of Specimens Collected," divided according to 
their habitats, rather than in a systematic list. The main vegetation regions of the 
country and the particular vegetation at each camp are described and discussed 
in relation to the distribution of the species throughout Africa and with regard to 
their possible bearing on the progress of speciation. Additional notes have been 
made on breeding seasons and on field habits and systematics where these have 
proved of particular interest. The usefulness of these discussions is necessarily 
limited by the lack of personal knowledge of the country outside the route of the 
expedition and the paper as a whole is intended only to supplement the work that 
is being done concurrently, chiefly in the United States, on other collections from 
Angola, so that finally there may be a real understanding of the place of the Angola 
avifauna in African ornithology. 

A cknowledgments 

For the expedition I was fortunate to have the support of the Zoology Depart- 
ment, British Museum (Natural History), and in being able to get together an Anglo- 
American party following the Pan- African Ornithological Congress in Livingstone 
in July 1957. The American members were Mr. and Mrs. Donald S. McChesney, 
Research Associates of Cornell University. Their chief interest was in photography 
and sound-recording of bird songs, although they took an active interest in the 
specimens collected during the time they were with the rest of the expedition : 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 371 

this time was unfortunately very limited as private affairs compelled them to return 
to the States on 2nd September. Mr. John Williams, from the Coryndon Museum, 
Nairobi, came to take charge of the skinning ; he also had to leave about the same 
time having been unluckily dogged by ill-health throughout. The chief collectors 
were Lt.-Gen. Sir Gerald Lathbury, K.C.B., D.S.O., M.B.E., and Mr. A. L. (Tony) 
Archer from Nairobi, who brought his Land Rover and Owaka and Ali as mechanic 
and cook. John Williams and I also collected when time allowed from skinning. 
During their time in Angola Mr. McChesney engaged Mr. Mario Pirelli from Luanda 
as an interpreter and guide to the country. 

The trustees of the Percy Sladen and of the Godman Trusts made grants towards 
my travelling expenses and the British share of the expedition, and a grant was 
made from the Purchase Grant of the Museum. Further expenses were born by 
Mr. McChesney, General Lathbury and myself, Mr. McChesney also being responsible 
for the financing of the American share. The Committee of the Bird Exploration 
Fund contributed with the loan of a gun and the gift of some ammunition. 

In Livingstone Mr. Rudyerd Boulton gave valuable advice on the country and 
habitats of the birds which was of great assistance in planning our route. In Angola 
we received unfailing help from all Government officials with whom we came in 
contact, and also helpful advice on the birds of the area from Mr. R. Braun at Sa 
da Bandeira. 

Since returning to the Museum I have received great assistance in working out 
the collection from colleagues in the Bird Room and Mr. A. W. Exell, in the Botany 
Department, and through loans and letters from many ornithologists including 
Dr. Dean Amadon, Professor J. Berlioz, Mr. C. W. Benson, Mr. P. A. Clancey, 
Dr. H. Friedmann, Mr. J. C. Greenway, Mr. R. E. Moreau, Prof. N. E. Collias, 
Miss M. Patterson, Captain C. R. S. Pitman, Dr. A. L. Rand, Prof. H. Schouteden, 
Dr. W. Serle, Mr. R. H. N. Smithers, Prof. E. Stresemann, Mr. M. P. Stuart Irwin, 
Mr. M. A. Traylor, and Mr. C. M. N. White. Mr. Smithers also kindly had some birds 
skinned for me in Bulawayo which had been preserved in salt. Mr. B. C. Lack, 
of the Metereological Office, supplied rainfall figures. Mr. C. W. Mackworth-Praed 
and the late Captain C. H. B. Grant have allowed me full use of the manuscript 
of the second part of the African Handbook of Birds covering Southern Africa. 
Dr. G. Rudebeck has supplied me with a list of birds collected on the Visser- 
Transvaal Museum Expedition to Angola in 1956, and allowed me to examine 
specimens. Commander A. M. Hughes and Mrs. M. Gull have drawn the maps. 

To all these people I am very much indebted, but above all to my companions 
for their contributions in their respective spheres, especially to the two collectors 
who were responsible for the unusually large variety of birds collected in two months. 

Deposition of Specimens Collected 

In all 980 specimens were obtained of 306 species, including the second and third 
known specimens of Prionops gabela. The majority were skinned on the spot, but 
some less important were preserved in salt for eventual treatment in the Museum 
(see Ibis, 1959 : 254), and fifty-four were preserved in spirit as anatomical specimens. 



372 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 



At the request of the Portuguese authorities a number of duplicate specimens have 
been sent to the Museum in Luanda. A proportion of the remainder, mainly from 
eastern districts, are in the American Museum of Natural History, a small number 
in the National Museum of Southern Rhodesia, Bulawayo, and the bulk in the 
British Museum. The photographs and sound-tracks have been deposited at 
Cornell University. 

ITINERARY 

The expedition entered Angola from the Belgian Congo along the line of the Ben- 
guela railway at Dilolo. It had been hoped to do some exploratory work in the little 
known areas of the east and south-east, but Williams' illness and shortage of time 
made this impossible. Thus little new ornithological ground was broken except 
in some parts along the main road from the frontier to Nova Lisboa, and energies 
were mostly concentrated on seeing as many different types of country as possible, 
and in filling the known gaps in the British Museum collection from such well- 
known districts as Mt. Moco, Sa da Bandeira, Chingoroi and Gabela. 

A full list of camps follows with the altitudes and dates so that these may be 
correlated with the Tables of Specimens collected at the different camps : — 



Date of 










arrival 


Locality of camp 


Lat. 


Long. 


Alt. 


27th J 


uly 


Luau R. Angola/Congo 
border 7 m. E. of Texeira 
de Souza 


10.42 S. 


22.12 E. 


• 3.600' 


1st August 


Luacano 


n. 16 S. 


21.38 E. . 


3.550' 


3rd 


,, 


Lake Dilolo 


11.30 S. 


22.00 E. 


3.500' 


6th 


,, 


Luacano 


— 


— 


— 


7th 


,, 


28 m. W. of Vila Luso 


11.47 s. 


19.52 E. 


4,400' 


8th 


>> 


5 m. W. of Munhango 


12.12 s. 


18.42 E. 


4,700' 


9th 


" 


5 m. W. of Vila General 
Machado 


12.03 s. 


17.30 E. 


4,800' 


10th 


a • 


Vouga 


12. II S. 


16.47 E. 


5.700' 


1 2th 


>> 


5 m. E. of Nova Lisboa 


12.44 s. 


15.47 E. 


5.5oo' 


14th 


„ 


Mt. Moco 


12.27 s. 


15.10 E. 


6,000- 
8,000' 


25th 


»» 


5 m. N. of Vila Flor 


12.41 s. 


I5-3I E. 


5.5oo' 


27 th 


>> 


Caconda 


1343 s. 


15.06 E. 


5,200' 


28th 


,, 


Quipungo 


14.49 s. 


14.34 E. 


4,400' 


29th 


,, 


Sa da Bandeira 


1455 s. 


13.30 E. 


5,600' 


30th 


>. 


Leba 


15.04 s. 


13.16 E. 


5,800' 


4th September . 


Cahinde (Tampa) 


15.30 s. 


13.27 E. 


. 2,500' 


6th 


,, 


5 m. S.W. of Sa da Ban- 
deira 


— 


— 


6,600' 


7th 


>, 


Chingoroi 


1337 s. 


14.01 E. 


3.300' 


nth 


$> 


12 m. S.E. of Benguela 


12.35 s. 


13.25 E. 


. S.L-500' 


13th 


" 


60 m. N. of Lobito on Novo 
Redondo road 


— 


— 


c. 800' 


14th 


„ 


20 m. N.E. of Novo Redon- 
do on Gabela road 


— 


— 


. c. 1,000' 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 373 



Date of 










arrival 


Locality of camp 


Lat. 


Long. 


Alt. 


15th 


. Cuvo R. 12 m. S.W. of . 
Gabela 


10,48 S. 


14.20 E. 


2,500' 


20th ,, 


40 m. S. (by road) of Mum- 
bondo 


10.09 s. 


14.15 E. 


1,000' 


25th 


. Calulo 


10.00 S. 


14.53 E. 


3,300' 


27th ,, 


Quimberima plantation 40 
m. S.E. of Calulo 


— 


— 


. 3»7oo' 



Other localities by roadside: 

9th August . Coemba, 40 m. W. of Mun- . — . — . — 

hango 
Saiangikilo R. 16 m. W. of . — . — . — 

Munhango 
13th ,, . Quipeio, 25 m. N.W. of . 12.26 S. . 1530 E. . c. 6,000' 

and Nova Lisboa 

25th 
7th September . Quilengues, on Sa da Ban- . 14.00 S. . 14.00 E. . c. 3,900' 
deira-Chingoroi road 



VEGETATION 

Main Vegetational Regions 

All the main vegetation regions of tropical Africa are represented in Angola with 
the exception of extensive marshlands of the " Sudd " type. These regions are 
indicated on the following map which has been adapted from the Vegetation Map 
of Africa published on behalf of 1' Association pour TEtude Taxonomique de la 
Flore d'Afrique Tropicale with the assistance of U.N.E.S.C.O. (Oxford University 
Press : 1959), with slightly more detail in those districts covered by the expedition. 
The regions can be summarized as follows : — 

(a) Savannah Woodland 

In eastern and central Angola this is largely of the Brachystegia and Isoberlinia 
type and is extensive, being found even up to 7,000 feet on the mountain slopes. 
For convenience it will be referred to throughout as Brachystegia woodland. 

In the west, in a narrow strip along and below the escarpment, there is rather 
richer vegetation, shown on the map as relatively moist types of savannah woodland. 
In this strip Brachystegia no longer predominates, there is more variety and Baobabs 
are common. 

(b) Grassland 

Patches of grassland often very flat and poorly drained are interspersed through- 
out the Brachystegia woodland. In the west there is more open country with many 
wide stretches of rolling grassland in which other vegetation is confinen to thickets 
and clumps, mostly in the river and stream valleys. 



374 THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 

Vegetation map of Angola 




Miles 

KEY TO VEGETATION MAP 



Montane Forest 



Woodlands and Savannahs with 
abundant Brachystegia 



Escarpment or Riverine Forest 



+ + + Grass Steppe, Luanda type 



VM 



Mixed Woodlands and Savannahs 
relatively moist types 



Semi-deser. 



Mixed Woodlands and Savannahs, 
relatively dry types with abun- 
dant Acacia 



Desert 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 375 

(c) Semi-desert and Thorn 

In the south and south-west the acacia thornveld of the Kalahari and South 
West Africa extends into Angola and along the coastal strip to the latitude of Lobito. 
Here the sandy plain gives way to the rolling grassland and the predominance of 
acacia diminishes northwards to Luanda. 

(d) Desert 

The Namib desert of South West Africa extends in a narrow strip up the coastal 
plain into southern Angola, merging into the semi-desert or scrub a little north of 
Mocamedes and in the foothills towards the base of the Chela escarpment. 

(e) Forest 

The forests of Angola, excluding Cabinda in which true tropical rain forest is 
found, are of three types. True montane forest of great antiquity is found in pockets 
on the highest mountains of the Bailundo highlands such as Mt. Moco and Mt. 
Soque. It was also fairly extensive on the Mombolo plateau when Boulton collected 
there in 1931, but we were informed in Nova Lisboa that it has largely been cut 
down. Remnants of forest on the Chela escarpment may also prove to be relic 
montane type. 

The lowland forests are of more recent origin and can be divided into what might 
be termed escarpment forest, and gallery or riverine forest. The escarpment forest 
is found between 1,000 feet and 3,000 feet in sometimes extensive patches below or 
on the face of the escarpment in areas which combine the high summer rainfall 
of inland districts with high all-the-year-round humidity and frequent mists due 
to the meeting of the cold waters of the Benguela current with the warm equatorial 
waters. 

Gallery forest is found in patches along the rivers of the north and east, and also 
along some of the west coast rivers below the escarpment. 

Vegetation at Camps 

The expedition traversed all the main vegetation regions with the exception of 
the true desert and it seems useful to discuss in more detail the country at each 
camp, and on the road between. The camps and districts discussed have been 
numbered to correspond with the columns in the Tables of Specimens collected 
(pages 379-39 1 )- 

(1) Luau River 

Most of the collecting was done in the narrow strip of riverine forest or the Brachy- 
stegia woodland around. There was no open grassland but some clearings in the 
woodland and the edges of cultivation. 

(2) Luacano Area 

There was some riverine forest and Brachystegia woodland, but the camp was 
chiefly notable for the wide flat treeless plain crossed by damp ditches which extended 
some miles east and south of Luacano village. 



376 THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 

(3) Lake Dilolo 

Most of the collecting was done in the grassy swamps and reed-beds on the border 
of the lake near the Government Rest House, and a little in some woodland a few 
miles to the west. There was no time to explore the extensive possibilities of the lake, 
the largest in Angola, or to organize the use of native dugout canoes which would 
be the most suitable transport for the purpose. 

(4) Roadside Localities between Luacano and Vouga 

All camps were made in Brachystegia woodland but this was interspersed with 
grassy dambos similar to the grassland at Luacano in their flatness and lack of 
bushes, but not so extensive. (Plate 5, b). 

(5) Vouga 

The grasslands here differed in being extensive undulating plains with moist 
marshy patches in the valleys and some patches of scrubby bushes. A small patch 
of stunted woodland with some bushes and cultivation surrounded the camp. 

(6) Nova Lisboa area 

This area was predominantly Brachystegia woodland with some grassy clearings 
and cultivation. 

(7) Foothills of Mt. Moco 

The lower slopes were covered with Brachystegia woodland with thick tangled 
grasses in the stream beds, and grassland and cultivation below. 

(8) Mt. Moco, over 6,000 feet 

Brachystegia was found up to 7,000 feet on some slopes and montane forest 
was limited to patches lining precipitous gullies at the head of the streams on the 
upper slopes, descending to about 6,800 feet on some of the eastern and southern 
slopes. The forest trees were nowhere very tall or very dense but mostly of ever- 
green species totally different from the woodland types and with tangled under- 
growth beneath. The bare slopes above the woodland and away from the forest were 
rocky and grassy with scattered aloes and small scrub, with numerous plants in flower. 

(9) Vila Flor, and localities on Luimbale-Nova Lisboa road 

This area was predominantly Brachystegia woodland with some grassy clearings 
and cultivation. 

(10) Caconda and Caconda-Quipungo road 

This country was similar to Vila Flor until a change was noted about 40 miles 
south of Caconda where the trees appeared much the same size and form but were 
notably barer of leaves. This change would not have been worthy of note except 
that it corresponded with some change in the bird life, four species being met within 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 377 

a few miles which had not hitherto been encountered, namely, Neocichla gutturalis, 
Thamnolea arnotti, Turdoides jardinei and Eurocephalus anguitimens. 

(11) Quipungo 

The first patch of thorn country was encountered here, with some grassy clearings 
and a small patch of Brachystegia. 

(12) Sa da Bandeira and the roads to Quipungo and Chingoroi 

This was very mixed country with thorn and Brachystegia alternating, and with 
richer and more varied vegetation northwards towards Chingoroi as the road 
descends to 3,000 feet. An overnight camp at Sa da Bandeira was made on top 
of the escarpment above the town and some collecting was done in the stunted 
Brachystegia on the slopes and the rocks above. 

(13) Leba and Humpata-Jau road 

The camp at Leba was in an area of mixed vegetation. It was situated about a 
mile inland from the edge of the Chela escarpment at a point where a stream cuts 
its way through a gorge in a series of waterfalls down 3,000 feet to the foothills and 
plains at Capangombe. Immediately around the camp was an area of tangled bush 
and thorn full of aromatic shrubs ; in some clearings by the river were remnants 
of forest and a patch of forest was found by the river near the foot of the escarpment. 

The rocks and fringing trees of the escarpment were the typical habitat of Xeno- 
copyschus ansorgei, and in one place along the edge was an extensive patch of tangled 
evergreen bush, difficult to penetrate, which contained few birds. Other slopes 
above the river were covered in stunted Brachystegia, and on the plateau running 
back from the escarpment patches of Brachystegia were mixed with patches of 
thorn and occasional grassy or cultivated clearings. (Plate 5, a). 

In such mixed country it was not always easy to define the habitat of specimens 
collected and this should be borne in mind when consulting the Tables. 

Some collecting of larks and pipits was done on open grassland on the Humpata-Jau 
road, and a party of water-birds was seen at a small lagoon in the centre of these 
grasslands. 

(14) Cahinde area 

Included in this area, is the road down the escarpment where the slopes are lined 
with stunted Brachystegia and Mopane. This gives way at the bottom of the escarp- 
ment among the foothills to typical semi-desert country of mixed thorn and Terminalia 
or Combretum, with the Mopane becoming less common westwards towards the desert. 
An excursion was made westwards for fifty miles from Cahinde without coming out 
of the foothills into true desert. 

(15) Chingoroi 

The patch of forest in which most of the collecting was done might be described 
as part riverine and part escarpment being associated with a stream, but more 



378 THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 

extensive than the usual riverine strips. Other birds were collected in reeds and grasses 
bordering the stream. 

(16) Benguela area (3-15 Miles South-east) 

The descent from Chingoroi to the coastal strip behind Benguela was gradual and 
through mixed country which gave way on the coastal plain to typical thorn bush 
resembling the Kalahari and the Vornamib. It was more open than the country 
at Cahinde with more grassland and acacia and no Mopane, and only broken by 
rocky kopjes. 

(17) Lobito — Novo Redondo-Gabela road 

The road to Novo Redondo wound over the rolling grasslands mentioned in the 
discussion on vegetational regions. Thorn bushes and Euphorbia were found on 
the hill-sides and frequent clumps of dry tangled thickets similar to those found on 
above the escarpment at Leba. Baobabs were common. Between Novo Redondo 
and the foot of the escarpment there were clumps of richer vegetation bordering 
streams and lagoons, but no collecting was done here. 

(18) Cuvo River, 12 Miles SW. of Gabela 

All collecting from this camp was done within three or four miles of the river and 
in the forest on the face of the escarpment, which is again precipitous at this latitude, 
though not to such a degree as at Leba. The forest is largely underplanted with 
coffee and the original undergrowth destroyed, but there was an uncleared patch in 
which we collected not far from the camp. The forest ended abruptly at the top of 
the escarpment before the town of Gabela and also a few miles further south along 
the road beyond our camp. The few species listed as collected in grassland in this 
camp were from tangled grass near the river's edge or a patch of disused cultivation 
in a forest clearing. 

(19) 40 Miles South of Mumbondo and road to Luanda 

Very little collecting was done on the road from Gabela to Luanda via Mumbondo 
and Muxima, but it was noteworthy for finding Prionops gabela well outside the 
forest area at the foot of the escarpment in a patch of mixed bush and woodland 
between cultivated clearings. 

(20) Calulo and Quimberima (40 Miles SE. of Calulo) 

After the main expedition was over I collected a few birds at Calulo which is 
situated in open grassland with some patches of stunted Brachystegia and bush, 
and on the plantation of Mr. and Mrs. O. Kroel at Quimberima, where they very 
kindly put me up for two nights. The Quimberima birds were collected in a tiny 
patch of forest edging a stream and bordered by Brachystegia. 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 379 

Photographs 

Photographs and further descriptions of many of the localities and types of 
vegetation discussed will be found in Lynes' two papers on his expeditions to Angola 
(1933 and 1938), in Heinrich's recent paper (1958), and in Gosswelier & Mendonca 

(1939)- 

TABLES OF SPECIMENS COLLECTED 
Key to Tables 

Species which appear in more than one table, or which are referred to in the Systematic and 
Field Notes, have cross references in the left hand columns. 

Collecting localities are grouped into districts for estimate of breeding dates and numbered as 
follows : 

1. Luau R. 

2. Luacano area : Texeira de Sousa — Luacano road. 

3. Lake Dilolo. 

4. Vila Luso — Vouga road. 

5. Vouga. 

6. Nova Lisboa area. 

7. Foothills of Mt. Moco (6,000 feet). 

8. Mt. Moco over 6,000 feet. 

9. Vila Flor : Quipeio : Nova Lisboa — Luimbale road. 

10. Caconda : Caconda — Quipungo road. 

11. Quipungo. 

12. Sa da Bandeira : Sa da Bandeira — Quipungo road : Sa da Bandeira — Chingoroi road. 

13. Leba : Humpata — Jau road. 

14. Cahinde : road down escarpment : Cahinde — Mocamedes road (40 miles). 

15. Chingoroi. 

16. Area 3 — 12 miles S.E. of Benguela. 

17. Lobito — Novo Redondo — Gabela road. 

18. Cuvo R., 12 miles S.E. of Gabela. 

19. Forty miles south of Mumbondo. 

20. Calulo : Quimberima (40 miles S.E. of Calulo). 
Unsexed specimens are listed by number only (i.e. 1 $, 2 $, 1). 
Specimens in spirit are listed in brackets. 

Young birds not fully grown are listed as " juv." 

Young birds fully grown are listed as " imm." 

" Summer " breeding season indicates any month between November and May. 



3 8o THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 

Table I. 



-Birds 





Other 


















references 




Eastern Districts 








Central 




Tables 


Syst. 
Notes 


1 


2 


3 


Breeding 


4 


5 


6 


4 Francolinus finschi 


— 


* 


— 


— 

















6 coqui angolensis 


— 


— 


— 


— 


— 


— 


— 


— 


16* 


15 Chrysococcyx klaasi . 


— 


— 


— 


(1?) 


— 


— 


— 


— 




16 Cuculus c. cafer 


— 


— 


— 


— 


— 


— 


1 6 


— 


— 


19 Poicephalus meyeri damarensis . 


II 


— 


— 


— 


— 


— 




— 


— 


24 Coracias spatulatus . 


— 


— 


— 


16 


— 


Summer 


— 


— 


— 


25 Halcyon chelicuti 


— 


— 


— 


(1) 


— 


— 


— 


— 


— 


28 Merops nubicoides 


— 


— 


— 


16* 


— 


? Sept. 


— 


— 


— 


29 Melittophagus pusillus meridionalis 


— 


— 


— 


— 


— 


— 


— 


— 


— 


30 Dicrocercus h. hirundineus . 


— 


— 


10* 


— 


(1) 


Summer 


— 


— 


— 


31 Tockus p. pallidirostris 


— 


— 


16* 


— 




— 


— 


— 


— 


34 Glaucidium perlatum licua . 


— 


— 


— 


— 


— 


— 


— 


— 


— 


37 Semeiophorus vexillarius 


— 


— 


— 


— 


— 


— 


— 


1 6 


— 


40 Caprimulgus pectoralis fervidus . 


— 


— 


3? 


— 


— 


Summer 


— 




— 


45 Colius striatus congicus 


— 


* 


id* 


— 


— 


? Aug. 


— 


— 


— 


47 castanotus 


II, IV 


* 


— 


— 


— 


— 


— 


— 


— 


48 Lybius torquatus bocagei . 


— 


* 


1 6*. 1? 


— 





? July onwards 











49 Buccanodon anchietae anchietae . 


— 


* 


— 


— 


— 


— 


— 


— 


— 


katangae . 


— 





_ 


_ 


16 


Summer 


1 6, 1 ? 








52 Tricholaema diadematum fronta- 


— 


— 


— 


— 


— 


— 


— 


— 


— 


tum 




















54 Pogonolius chrysoconus extoni 


IV 


— 


— 


— 


2? 


? Aug. /Sept. 


— 


— 


— 


56 Prodotiscus regulus . 


— 


— 


1 6*. 1 ¥ 


— 


— 


— 


— 


— 


— 


57 insignis lathburyi 


— 


— 


— 


— 


— 


— 


— 


— 


— 


59 Campethera cailliautii fulleborni 


— 


* 


id* 


— 


— 


? June 


1 ? 


— 


— 


5 permista 




















61 Dendropicos fuscescens hartlaubii 


IV 


* 


16* 


1? 


— 


? Sept. onwards 


— 


— 


— 


62 Mesopicos griseocephalus persimilis 




* 


— 


— 


— 


— 


— 


— 


— 


76 Anthus similis schoutedeni . 


— 


* 


— 


— 


— 


— 


— 


— 


— 


yy lineiventris 


— 


* 


— 


— 


— 


— 


— 


— 


— 


82 Turdoides jardinei tamalakanei . 


— 


— 


— 


— 


— 


— 


— 


— 


— 


94 Alseonax adustus angolensis 


— 


— 


— 


— 


— 


— 


1 6", 1 ? 


— 


— 


angolensis % subadustus 


— 


— 


i? 


— 


— 


? Sept. 


— 


— 


— 


97 Myopomis bohmi 


— 


— 


— 


— 


— 


— 


1 6" 


— 


— 


101 Bradornis pallidus murinus 


IV 


— 


— 


— 


— 


— 


16", 1? 


— 


— 


103 Melaenornis pammelaina 


— 


— 


— 


— 


— 


— 


i? 


— 


— 


105 Hyliota australis slatini (?) . 


— 


* 


— 


— 


— 


— 


3 6", 1? 


— 


— 


106 flavigaster barbozae 


Ill 


— 


— 


— 


— 


— 


16" 


— 


— 


107 Batis m. molitor 


— 


— 


16" 


(2?) 


(1 6), 1 ? 


? Sept. 


— 


— 


— 


in Platysteira peltata mentalis 


II 


— 


— 


I $ 




Summer 


— 


— 


— 


113 Elminia albicauda 


II 


— 


— 


I? 


— 


— 


— 


— 


— 


114 Tchitrea viridis plumbeiceps 


II 


* 


— 




— 


— 


— 


16*. 1? 


— 


116 Turdus libonyanus verreauxi 


— 


— 


16* 





— 


? Sept. 


— 


— 


— 


119 Geokichla litsipsirupa stierlingi . 


— 


* 


— 





— 


— 


1? 


— 


— 


120 Monticola angolensis . 


— 


— 


— 





— 


— 


— 


— 


— 


124 Cercomela familiaris falkensteini . 


IV 


— 


— 





— 


— 


— 


— 


— 


125 Myrmecocichla arnotti harterti . 


— 


— 


— 





— 


— 


— 


— 


— 


137 Erythropygia b. barbata 


— 


— 


— 





— 


— 


1 6 


— 


— 


147 Calamonastes simplex huilae 


— 


* 


— 





— 


— 


— 


— 


— 


152 Sylvietta r. runcapilla 


— 


* 


— 





1? 


?Oct. 


— 


— 


— 


155 Eremomela icteropygialis polio- 


IV 


— 


— 





— 


— 


1 


1 


— 


xantha 




















156 Eremomela scotops pulchra 


— 


— 


— 


— 


1 6" (2) 


? Sept. 


16" 


— 


— 


157 atricollis . 


— 


— 


— 







— 


26 


— 


— 


158 salvadorii 


— 


* 


— 





— 


— 




— 


— 


159 Camaroptera brevicaudata subsp.? 


II, IV 


* 


1 6* (i) 





— 


— 


— 


— 


— 


162 Cisticola chiniana fortis 


IV 


— 


— 





— 


— 


— 


— 


— 


166 fulvicapilla dispar 


IV 


— 


— 


— 


— 


— 


— 


— 


— 


dispar % hallae 


— 


— 


— 





16" 


Summer 


— 


— 


— 


177 Prinia subflava graueri 


IV 


— 


— 





16" 


Summer 


— 


— 


— 


179 flavicans bihe . 


IV 


— 


— 







— 


— 


26*, 2? 


— 


191 Psalidoprocne orientalis reichenowi 


— 


— 


2 6* (1) 


1 imm. 6* 


— 


? June 


— 




— 


195 Coracina pectoralis 


— 


— 


1$ 


— 


— 


Summer 


— 


— 


— 


197 Prionops plumata poliocephala . 


— 


— 


— 


— 


16* 


? June 


1 


— 


— 


198 retzii nigricans 


— 


— 


— 


— 


— 


— 


16* 


— 


— 


200 Eurocephalus anguitimens . 


— 


— 


— 


— 


— 


— 


— 


— 


— 


201 Nilaus afer amnis 


— 


. — 


— 


— 


— 


— 


— 


— 


16* 


202 Lanius s. souzae 


— 


— 


— 


16* 


— 


Summer 


— 


16* 


16* 


205 Laniarius aethiopicus major 


— 


* 


1 imm, 6* 


16" 


— 


? May/June 


— 






208 Dryoscopus cubla hamatus . 


II, IV 


— 


— 


— 


— 


— 


16" 


— 


— 


211 Tchagra s. senegala . 


— 


* 


1 6", 1? 


1 ? 


— 


? June 


— 


— 


— 


213 Antichromus minutus anchietae . 


— 


— 


1 $ 


— 


— 


— 


— 


— 


— 


216 Chlorophoneus sulfureopectus 


II 


— 


— 


16", 1 


— 


— 


— 


— 


— 


similis 




















218 Malaconotus hypopyrrhus inter- 


— 


* 


1 6" 


— 


— 


? June 


1$ 


— 


— 


positus 




















222 Parus leucomelas insignis . 


IV 


— 


16" 


— 


— 


Summer 


— 


— 


— 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 

of the Woodlands 



381 



Districts 








M 


. Moco 


South-central and South-western Districts 


7 


9 


20 


Breeding 


8 


Breeding 


10 


II 


12 


13 


15 


Breeding 




— 


— 


— 


— 


id\ i9 


? Sept. 


— 


_ 


— 


— 


_ 


— 


4 


— 


— 


— 


? June 


— 


— 


— 


— 





— 


— 


— 


6 


— 


— 


- 


Summer 


- 


— 


- 


- 


ic? 


- 


— 


— 


15 
16 
19 

24 


- 


i9 


— 


Summer 


- 


- 


— 


— 


- 


- 


- 


- 


25 
28 
29 
30 


— 


— 


- 


Summer 


— 


- 


— 


- 


— 


1 9, 


— 


- 


31 
34 
37 
40 


1 <J, 2 ?, 





— 


July 





— 





— 








— 


— 


45 
47 


I JUV. (? 


























3c? 
Id 


- 


- 


July/Aug. 

? June 
? Sept. 


= 


= 


T 


— 


= 


= 


- 


= 


48 
49 


— 


— 


— 


— 


Ic? 


— 


— 


— 


— 


Summer 


52 


ic? 


2c? 


- 


?Oct. 


- 


- 


- 


- 


Ic? 


- 


- 


?July 


54 
56 
57 
59 


— 


— 


- 


- 


19 


? Sept. 

















i9 


_ 


Summer 








Ic? 





_ 








Summer 


61 


— 


— 


— 


— 


— 


— 


— 


— 


— 


1 imm. 


— 


— 


62 


1 


1 


— 


— 


— 


— 


I<? 


— 


— 


— 


— 


?Oct. 


76 


— 


— 








ic? 


Summer 














77 


— 


— 


— 


— 




— 


ic?, 19 


— 


— 


— 


— 


Summer 


82 


— 


— 


— 


— 


1? 


? Sept. 


2 


— 


ic? 


— 


— 


?Oct. 


94 





I(J 





? Sept. onwards 























— 


97 


l«J 




— 


? July onwards 
?July 
? Sept. 


ic? 


Summer 


— 


— 


— 


— 


— 


— 


101 
103 
105 



































1 c? 








? Sept. 








— 


— 











— 


106 


icJ, 1? 


ic? 


— 


? Sept. 


— 


— 


— 


— 


— 


— 


— 


— 


107 
in 


— 


— 


— 


? Aug. 


— 


— 


— 


— 


— 


— 


— 


— 


113 
114 


1? 


I <?, 19 


— 


? Sept. 

? Sept. 

? May /June 


— 


— 


— 


— 


— 


— 


— 


— 


116 
H9 
120 








1 imm. cj 


























— 


— 


— 


— 


id\i9 


? Sept. 


— 


— 


— 


— 


— 


— 


124 


— 


— 


— 


— 




— 


2C?, I 9 


— 


2c? 


— 


— 


? Sept. 


125 


— 


19 


— 


? Sept. onwards 


— 


— 




— 




— 


— 


— 


137 


1 9 


1 9 


— 


? Sept. 


— 


— 





— 


— 


— 


— 


— 


147 




It? 


— 


?Oct. 


— 


— 


I? 


— 


— 


— 


— 


?Oct. 


152 
155 


(1) 


IcJ (3) 




? Sept. 




_ 


_ 


_ 




_ 


_ 





156 




!<?, 19 


— 


? Sept. onwards 


— 


— 


19 


— 


— 


— 


— 


? Sept. 


157 

158 


— 


19 


— 


? Sept. 


— 


— 





— 


— 


— 


— 


— 


— 





— 


— 


— 


— 





— 


— 


— 


— 


— 


159 











Summer 


























162 


— 


2c? 


— 


Summer 


— 


— 


— 


- 


— 


— 


— 


— 


166 


icJ. I? 


(I) 





? July onwards 


1 imm. $ 


? May/June 














Ic? 


?July 


177 


— 




— 


? Sept. onwards 


— 


— 





— 


— 


— 


— 


— 


179 


Ic? 





— 


? June 


— 


— 





— 


— 


— 


— 


— 


191 
195 
197 
198 


— 


- 


- 


?Oct. 


- 


- 


- 


29 


— 


— 


I 


?July 


1 <?, 1 9 


i(5,i? 


— 


? Sept. 


— 


— 


ic? 


— 


— 


— 


— 


— 


200 
201 






— 


Summer 


ic? 


Summer 


— 


— 


z 


— 


— 





202 
205 


i9 


E 


- 


? June 


1 


E 


- 


- 


- 


- 


E 


- 


208 
211 
213 
216 

218 











? Sept. 

















, 








— 


— 


1 imm. 9 


? May /June 


— 


— 


— 


— 


— 


— 


— 


— 


222 



382 THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 

Table L- 





Other 


















references 




Eastern Districts 








Central 






Syst. 


















Tables 


Notes 


1 


2 


3 


Breeding 


4 


5 


6 


223 Parus r. rufiventris . 


— 


— 


— 


io* 


— 


? Sept. 


— 


— 


— 


224 Oriolus auratus notatus 


— 


— 


i<? 


— 


— 


? June 


1 6 


— 


— 


225 Neocichla g. gutturalis 


— 


— 


— 


— 


— 


— 


— 


— 


— 


226 Cinnyricinclus leucogaster ver- 


— 


— 


1 0* 


— 


— 


? June 


— 


— 


— 


reauxi 




















228 Lamprotornis mevesii benguelensis 


IV 


* 


— 


— 


— 


— 


— 


— 


— 


229 acuticaudus . 


II 


— 


— 


1 6* 


— 


? Aug. 


1 imm. 9 


— 


— 


231 Zosterops senegalensis quanzae . 


— 


* 


— 


— 


— 


— 


— 


— 


— 


heinrichi 


II 


— 


— 


— 


— 


— 


1 6* 


— 


— 


quanzae £ anderssoni 


— 


— 


— 


— 


— 


— 


— 


— 


— 


232 Nectarinia kilimensis gadowi 




* 














1 6, 1 9. 

2 imm. 6, 
1 imm. 


233 bocagei 


— 


* 


— 


— 


— 


— 


— 


— 


— 


235 Cinnyris cupreus chalceus . 


— 


— 


i? 


(1) 


16 


Summer 


1? 


— 


26(1) 


237 0. oustaleti . 


~~ 


# 


~~ 










1 6 


1 ?, 
1 imm. $ 


238 venustus falkensteini 


— 


— 


— 


26*, 
1 imm. 6* 


— 


? June onwards 


16 


— 


(2) 


239 afer ludovicensis . 


II 


— 


— 


— 


— 


— 


— 


— 


— 


240 chalybeus intermedius . 


— 


— 


— 


— 


— 


— 


(2) 


— 


— 


242 Chalcomitraamethystinademinuta 


— 


— 


2<J(I) 


— 


— 


? June 


36 


2? 


— 


243 senegalensis saturatior 


— 


— 


— 


16* 


16*, 1 $ 


? June 


16* 


— 


1 


246 Anthreptes collaris phillipsi 


II 


* 






1 imm. 6* 
1 imm. 9 


June 


1 6 




— 


247 longuemariae angolen- 


— 


— 


I<? 


— 


— 


? Sept. 


— 


— 


— 


248 anchieti . 


— 


— 


— 


— 


— 


— 


— 


— 


— 


249 Salpornis spilonota salvadori 


— 


— 


— 


— 


— 


— 


16* 


— 


— 


254 Petronia superciliaris . 


— 


— 


I? 


— 


16 


?SeP 


16* 


— 


— 


258 Ploceus x. xanthops . 


II 


— 


— 


— 


— 




— 


— 


— 


269 Lonchura cucullata scutata 


— 


— 


(I) 


— 


— 


— 


— 


— 


— 


274 Pytelia m. melba 


IV 


— 


— 


— 


— 


— 


— 


— 


16* 


289 Steganura paradisaea obtusa 


— 


— 


I? 


— 


— 


? Aug. 


16* 


— 


— 


291 Serinus sulphuratus sharpii 


— 


— 


— 


— 


— 


— 


— 


— 


1 


295 Poliospiza gularis benguellensis . 


— 


— 


— 


— 


— 


— 


— 


— 


— 


297 Emberiza cabanisi cognominata . 


— 


— 


— 


— 


— 


— 


16* 


— 


— 


298 f. flaviventris 


IV 




2 6*, 

1 imm. 6* 






? May /June 









Table II. — Birds 









Other references 


Eastern Districts (riverine) 




Tables 


Syst. 
Notes 


1 


2 


Breeding 


1 Accipiter minullus 





— 


— 








2 Francolinus swierstrai . 






— 


* 


— 


— 


— 


14 Turtur tympanistria fraseri . 






— 


— 


— 


— 





17 Ceuthmochares a. aereus 






— 


— 


— 


— 





19 Poicephalus meyeri matschei 






I 


— 


— 


16* 


— 


21 Tauraco corythaix schalowi . 






— 


* 


— 




— 


22 erythrolophus 






— 


* 


— 


— 


— 


27 Ispidina p. picta .... 






— 


— 


16* 


— 





36 Ciccaba woodfordii bohndorffi 






— 


— 


16* 


— 


Summer 


41 Caprimulgus poliocephalus koesteri 






— 


* 




— 


— 


43 Heterotrogon vittatus camarunensis 






— 


* 


— 


— 


— 


44 Apaloderma n. narina . 






— 


* 


— 


— 





47 Colius castanotus 






I, IV 


* 


— 


— 





50 Gymnobucco calvus vernayi 






— 


* 


— 


— 





53 Viridibucco coryphaea angolensis . 






— 


* 


— 


— 





55 Indicator m. minor % conirostris . 






— 


* 


16 


— 


? July 


58 Campethera abingoni annectens 






IV 


— 


i9 


— 


Summer 


60 nivosa herberti . 






— 


* 


— 








62 Mesopicos griseocephalus persimilis 






I 


* 


— 


— 





63 Smithomis capensis albigularis 






— 


— 


26*, i9 


— 





83 Pseudoalcippe abyssinicus ansorgei 






— 


— 




— 





84 Malacocincla fulvescens dilutior 






— 


« 











86 Pycnonotus barbatus tricolor 





















87 Andropadus 1. latirostris 





















88 virens zombensis 






— 


— 


26* 


1 6*, 1 9 


? June 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 



383 



Continued 



Districts 


Mt. Moco 


South-central and South-western Districts 


7 


9 


20 


Breeding 


8 


Breeding 


10 


11 


12 


13 


15 


Breeding 



































_ 


223 


— 


— 


— 


?Aug. 


— 


— 


— 


— 


— 


— 


— 





224 


— 


1 — 


— 


— 


— 


— 


3(?,2$(l) 


— 


— 


— 


— 


Summer 


225 


ic? 


— 


— 


?July 


— 


— 


— 


— 


— 


— 


— 


— 


226 








— 


— 


— 


— 


— 


— 


I? 


— 





Summer 


228 


I<J,1? 


— 


— 


? May/June 
and Sept. 


— 


— 





— 


— 


— 


— 


— 


229 


— 


1? 


z 


? Sept. 
?July 


1 








z 






— 


— 


231 


5 c?, 


ic?(i) 




June 


4 c?, 1? (1) 


PJuly 




z 


— 


Ic? 


— 


? July 


232 


1 imm. (J 


























- 


i<J, 1 9 
3 imm <j> 


— 


June 


— 


— 


— 


— 


— 


— 


— 


- 


233 


— 


(3) 


— 


Summer 


(1) 


— 





— 


— 


— 


— 


— 


235 


4 <?, I JUV. 


2<J, 


— 


June onwards 


— 


— 


— 


— 


— 


— 


— 


— 


237 


(I) 


2 imm. $ 


























1? 


— 


Summer 


— 


— 





~~ 


— 


Ic? 


— 


— 


238 


I«J 




- 


- 


- 


- 


i imm. <J. 

1? 


- 


I? 


- 


- 


- 


239 
240 


— 


iimm.<J(i) 


— 


June 


Ic? 


? June 














242 


2 


1 imm. (J 


— 


? June 


— 


— 


— 


— 


z 


z 








243 
246 


- 


1?, 
1 imm. 6* 


- 


- 


- 


- 


ic?, 
1 imm. 6* 


- 


- 


- 


- 


- 


247 


I? 


2$ 


— 


? June 


— 


— 


— 


— 


— 


— 


— 


— 


248 


2c? 




ic? 


PSept. 


— 


— 


— 


— 


Ic? 


— 


— 


PAug. 


249 


It? 


ic? 


— 


? Sept. 


— 


— 


— 


— 


— 


— 


— 


— 


254 


I<?, I? 


— 


— 


? Sept. 


1? 


? Sept. 


— 


— 


— 


— 


— 


— 


258 


I 


— 


— 


— 


(1) 


— 


— 


— 


— 


— 


— 


— 


269 





— 


— 


PJuly 


— 


— 


— 


— 


— 


— 


— 


— 


274 











? Aug. 


— 


— 




















289 


2c? 


— 


— 


? July onwards 


Ic? 


PJuly 
? Sept. 


— 


— 


— 


ic? 


— 


— 


291 





— 


— 


— 


I? 














295 


— 


— 


— 


Summer 


— 


— 


— 


— 


— 


— 


— 


— 


297 


" 






















" 


298 



of the Forests 



Mt. Moco (montane) 


South-weste 


rn (? montane) 


Western (escarpment and riverine) 


8 


Breeding 


13 


Breeding 


15 


18 


20 


Breeding 














Ic? 











1 


1 6*, 1 imm. 6* 
i<? 


? May /June 


— 


— 




1? 


— 


— 


2 
14 


X? 


? Sept. 


7? 





Ic? 

I 





POct. 


17 
19 
21 
22 
27 
36 
41 


i<J 


Summer 


— 


— 


— 


— 


— 


— 


ic?, I? 


? Sept. 


— 


— 


— ■ 


— 





— 


43 


— 


— 


— 


— 


— 


1? 





?Aug. 


44 


I? 


— 


— 


— 


ic? 


— 





PJuly 


47 


2? 


? Sept. 


— 


— 


— 


— 







50 


Ic?, I? 


z 




z 


z 


z 


z 


z 


53 
55 





— 


z 





z 


ic? 





Summer 


58 
60 


1 6\ 2 imm. 


— 


— 


— 


— 


— 


— 


— 


62 


— - 


— 


— 


— 


1 imm. 6* 


— 


— 


July 


63 


3 c?, 1? 


? Aug. onwards 


— 


— 


— 


— 


— 




83 


— 


— 


— 


— 


— 


3<? 


— 


PJuly 


84 


Ic? 


PJuly 













— 




86 






— 


' — 


— 


!<?, I? 


— 


PJuly 


87 


— 


— 


— 


— 


— 





— 


— 


88 



ZOOL. 6, 7. 



23 



384 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 



Table II. — 



Other references 



Tables 



Eastern Districts (riverine) 



Syst. 
Notes 



Breeding 



89 Chlorocichla f . falkensteini . 

90 flaviventris occidentalis 

91 flavicollis flavigula 

92 Phyllastrephus fischeri cabanisi 

93 fulviventris . 

95 Alseonax cassini .... 

98 Chloropeta natalensis major . 

99 Parisoma plumbeum orientale 

104 Dioptrornis brunneus bailundensis. 

109 Batis m. margaritae 

no minulla .... 

in Platysteira peltata mentalis . 

112 Dyaphorophyia concreta ansorgei . 



113 Elminia albicauda 

114 Tchitrea viridis plumbeiceps . 

melampyra bannermani 

117 Turdus olivaceus stormsi 

118 pelios bocagei . 

131 Cossypha heuglini heuglini . 

subrufescens 

132 polioptera polioptera 

133 natalensis intensa . 

larischi . 



134 b. bocagei . 

135 Alethe p. poliocephala . 

136 Cichladusa ruficauda . 
139 Erythropygia leucosticta reichenowi 

142 Seicercus laurae .... 

143 Bradypterus mariae boultoni 

149 Apalis rufogularis brauni 

150 cinerea grandis . 
153 Sylvietta virens tando . 
159 Camaroptera brevicaudata sharpei (?) 

harterti 
169 Cisticola erythrops lepe 
178 Prinia 1. leucopogon 
180 Hirundo abyssinica unitatis 

193 Campephaga phoenicea sulphurata 

194 q. quiscalina 
196 Dicrurus 1. ludwigii 
199 Prionops gabela . 
203 Lanius mackinnoni 

208 Dryoscopus cubla hamatus 

209 a. angolensis 

214 Chlorophoneus multicolor batesi . 

215 nigrifrons manningi 

216 su&ureopectus similis 

217 Telephorus viridis 
219 Nicator vireo tando 
229 Lamprotornis acuticaudus . 
231 Zosterops senegalensis quanzae £ kasaica 
234 Cinnyris superbus 
236 bifasciatus 
239 afer ludovicensis 



244 Cyanomitra verticalis bannermani 

245 olivacea cephaelis 

246 Anthreptes collaris somereni . 

phillipsi . 
252 Passer griseus griseus . 

258 Ploceus x. xanthops 

259 Symplectes bicolor kigomaensis 

amaurocephalus 

260 Hyphanturgus n. nigricollis . 

261 ocularis crocatus . 

270 Spermaphaga r. runcapilla . 

271 Cryptospiza r. reichenovii 

273 Hypargos niveoguttatus 



285 Estrilda p. perreini 

288 Hypochaera nigerrima . 

296 Poliospiza burtoni tanganjicae 



IV 



IV 
I, IV 



16" 



i6\i? 



16*, 1? 



16* 

1 6*1 1 imm. 
1(2) 



26* 



i?_(i) 
1 imm. $ 



? June 
May/June 



? Sept. 
? June 

? May/June 



6*, 1? 



1 imm. 6* 



May/June 



16* 



io\ i? 

16* 
1 imm. o* 



1 6*, 1 imm. 6*, 
1 imm. $ 



1 $, 1 (1) 



? June 

Summer 
? May/June 

? May /June 
? June 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 



385 



Continued 



Mt. Moco (montane) 


South-western (? montane) 


Western (escarpment and riverine) 




8 


Breeding 


13 


1 
Breeding 


15 


18 


20 


Breeding 




- 


- 


- 


- 


16* 
16*, i? 
1 imm. $ 


1 ?, 1 imm. $ 


- 


? June/July 


89 
90 
















i<J 


? June 


— 


— 


— 


— 


— 


— 


9i 


3 6\ 2? 


Summer 


— 


— 


— 


— 


— 


— 


92 





— 





_ 


1 6*. 1?, 1 


16*, i9 


— 


? Aug. 


93 


— 


— 


— 


— 


— 


1 9 


— 


? Sept. 


95 
98 
99 


1_ 











16* 


— 





?Oct. 


3 6", 2$ 


? Sept. 


— 


— 




— 


— 


— 


104 


2 6*. I? 


? July onwards 


— 


— 


— 


— 


— 


— 


109 


— 


— 


— 


— 


16*, 2? 


16*, i 9 


— 


? Aug. 


no 


2 6*. 2? 


— 


— 


— 


— 


— 


1? 


?Oct. 


hi 




— 


— 


— 


3 6*, 3 9, 
1 juv. 6* 


16*, 1 9 




?July 


112 


16* 


— 


— 


— 


— 


— 


1? 


?July 


"3 


id", 1? 


? Sept. 


— 


— 


— 


— 


— 


— 


114 


— 


— 


— 


— 


— 


1? 


— 


?July 




— 


— 


— 


— 


— 


— 


— 


— 


117 


16* 


? Sept. 


— 


— 


— 


— 


— 


— 


118 


— 


— 


— 


— 


— 


— 


— 


— 


131 


16* 


?Oct. 


i6\ i? 


— 


1? 


— 


— 


Summer 




— 


— 


— 


— 


— 


— 


— 


— 


132 


— 


— 


— 


— 


— 


— 


— 


— 


133 


— 


— 


— 


— 


— 


1 6*, 2 ?, 
1 imm. $ 


— 


July 




6 6*. 2? 


Summer 


i? 


Summer 


— 


— 


— 


— 


134 




— 




— 


— 


1 juv. <J 


— 


Aug. 


i35 


— 


— 


— 


— 


16*, 3 9 


— 


— 


Summer 


136 


— 


— 


— 


— 


1 imm. $ 


— 


— 


? June 


139 


2 6*, 2 9, 2 (1) 


August 


— 


— 


— 


— 


— 


— 


142 


1 6", 2? 


? Sept. onwards 


— 


— 


— 


— 


— 


— 


143 


— 


— 


— 


— 


— 


1 6*, 2 juv. ? 


— 


Aug. 


149 


2 6", 2? 


? Sept. 


— 


— 


— 


— 


— 


— 


150 


— 


— 


— 


— 


— 


J 6, 1 9 


— 


— 


i?3 


— 


— 


16* 


— 


1 <J, 1 juv. 




— 


Aug. 


159 


■ — 


— 


— 


— 


— 


1 9 


— 


?July 




io\ 3? 


Summer 


— 


— 


i9 




— 


? July 


169 
178 
180 


1 


? Aug. 


— 


— 


— 





— 





— 


— 


— 


— 


— 


1? 


— 


?July 


193 


— 


— 


— 


— 


16* 




— 


?July 


194 


— 


— 


— 


— 


16", i9 


— 


— 


?Oct. 


196 


— 


— 


— 


— 




1 imm. $ 


— 


? June/July 


199 


— 


— 


— 


— 


— 


1 imm. 6* 


— 


? June /July 


203 


— 


— 


— 


— 


1 imm. ? 


26* 


— 


? June 'July, 
?Oct. 
?July 


2C* 


— 


— 


— 


— 


— 


16*, 1 9 


— 


20J 


— 


— 


— 


— 


— 


1 6*, 1 imm. 6* 


— 


? June/July 


214 


— 


— 


_ 


— 


1 imm. ?, 1 





— 


? June /July 


215 
216 


— 


— 


— 


— 


16*, i9 


— 


— 


— 


217 


— 


— 


— 


— 




I? 


— 


?July 


219 


1 9, 1 


— 


— 


— 


— 


— 


— 


— 


229 


— 


— 


— 


— 


— 


1? 


— 


?July 


231 


— 


— 


— 


— 


1 6* 


— 


— 


— 


234 


— 


— 


— 


— 


1 imm. o\ 1 9 


— 


— 


— 


236 


2 6\ 2 ?, 


?July 


2 6*. 1 ?, 


?July 


— 


— 


— 


— 


239 


5 imm. <J, 




2 imm. 6* 














4 imm. $ 


















— 


— 


— 


— 


— 


2(J,t imm. 6* 


— 


— 


244 
245 


— 


— 


— 


— 


— 


16, 1 9 


— 


— 


246 


— 


— 


— 


_ 


— 


16* 


— 


?July 


252 


— 


— 


26" 


— 


— 


i9 


— 


— 


258 
259 


— 


— 


— 





2 9, 1 imm. 6* 


1? 


— 


? June/July 


— 


— 


— 


— 


— 


2 imm. ? 


— 


? June /July 


260 


1 6\ 1 imm. 6* 


June 


— 


— 


— 


— 


— 


— 1 


26l 


— 


— 


— 


— 


— 


1 6*, 1 imm. ? 


— 


?July 


270 


— 


— 


— 


— 


— 


1 6*, 1 9, 


— 


July 


271 

















1 imm. 6* 








273 














i9 







?July 


285 


— 


— 


— 


— 


1 imm. 


— 


— 


July 


288 


1 6", 2? 


? Sept. 


— — 


— 


— 


— 




296 



386 THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 

Table III. — Birds of Grasslands, 





Other 


















references 




Eastern Districts 






Central 






Syst. 


















Tables 


Notes 


1 


2 


3 


Breeding 


4 


5 


6 


5 Francolinus levaillantoides jugularis 


— 


— 


— 


— 


— 


— 


— 


— 


— 


7 albogularis meinertzhageni . 


— 


* 


— 


2 6,39 


— 


Summer 


— 


— 


— 


8 Pternistes afer crainchii 


IV 


— 


— 


— 


16" 


? June 


— 


— 


— 


io Glareola pratincola fulleborni 


— 


— 


— 


— 


1? 


Summer 


— 


— 


— 


12 Cursorius temminckii .... 


— 


— 


— 


— 


— 


— 


1 juv. 6* 


— 


— 


1 8 Centropus toulou grilli .... 


— 


— 


— 


— 


— 


— 




— 


— 


26 Halcyon albiventris .... 


— 


— 


— 


I 


— 


— 


— 


— 


— 


38 Caprimulgus natalensis mpasa (?) . 


— 


* 


— 


— 


16" 


Summer 


— 


— 


— 


39 fossii .... 


— 


— 


— 


— 


2 6, 2 9 


Aug. onwards 


— 


— 


— 


42 Micropus caffer ansorgei 


— 


— 


— 


— 




— 


i<J 


— 


— 


64 Mirafra africana occidentalis 


— 


* 


— 


— 


— 


— 


— 


— 


— 


occidental is % pallida 


— 


— 


— 


— 











, — 


— 


kabalii 


— 


— 


— 


2 6*, 2? 


— 


Summer 


— 


— 


. — 


65 angolensis angolensis . 


— 


* 


— 


— 


— 


— 


1? 


3 6,i9 


— 


antonii 


— 


— 


— 


6 6\3? 


— 


Summer 


— 


— 


— 


66 rufocinnamomea fischeri 


— 


— 


— 


1? 


— 


— 


16 


— 


— 


69 Certhilauda albofasciata obscurata . 


— 


— 


— 




— 


— 




3 6,29 


— 


70 Calandrella cinerea saturatior 


— 


— 


— 


2<J, 2? 
2 JUV. 


— 


July 


— 




— 


72 Eremopterix verticalis damarensis . 


IV 


— 


— 





— 


— 


— 


— 


— 


73 Motacilla capensis simplicissima 


— ' 


— 


— 





— 


— 


— 


— 


— 


74 Anthus vaalensis neumanni . 


— 


* 


— 


I? 


1$ 


Summer 


i? 


2 6,i9 


i9 


75 leucophrys bohndorni 


— 


* 


— 






— 






— 


78 Macronyx ameliae .... 


— 


* 


— 


1 6, 2 ?, 
1 imm. 9 


16 


? July onwards 


— 


26 


— 


79 grimwoodi .... 


— 


* 


— 


i<? 


— 


?Oct. 


— 


2 6,i9 


— 


80 fulleborni ascensi . 


— 


* 


— 


2? 


io* 


?Oct. 


— 


i6,i9,i 


— 


81 Turdoides leucopygia hartlaubii 


IV 


— 


— 




1 imm. 9 


? May/June 


— 




— 


106 Hyliota flavigaster barbozae . 


I 


— 


— 





— 


— 


— 


— 


— 


122 Oenanthe pileata livingstonii (?) 


— 


— 


— 





— 


— 


— 


16 


— 


126 Myrmecocichla nigra .... 


— 


— 


— 


io\ 1? 


— 


Summer 


— 




— 


127 tholloni .... 


— 


* 


— 




— 


— 


— 


26,19 


— 


128 Saxicola torquata caffra 




— 


— 


2<J 


— 


— 


— 


1 6, 1 9, 


— 


141 Acrocephalus baeticatus cinnamomeus 


— 


* 


— 














eggs 





145 Calamocichla gracilirostris winterbottomi 


— 


* 


— 


— 


2 6,2? 


? Sept. 


— 


— 


— 


cunenensis 


— 


— 


— 


— 


— 


— 


— 


— 


— 


148 Schoenicola brevirostris alexinae 


— 


— 


1 imm. 9 


— 











— 


— 


160 Cisticola a. ayresii .... 


— 


— 


— 




















161 lateralis vincenti 


— 


— 


3 6\ 2? 


— 


— 


Summer 


— 


— 


— 


162 natalensis katangae . 


— 


— 


(1) 


— 


2? 


Summer 


— 


— 


— 


huambo . 


— 


— 


— 


— 


— 


— 


— 


— 


— 


164 tinniens perpulla 


— 


— 


— 


16 


I,?, I $ 


Summer 


— 


26 


— 


165 robusta angolensis . 


— 


— 


— 




I? 


Summer 


— 


16 


— 


167 brachyptera loanda . 


— 


— 


1? 


— 


16 


Summer 


16* 




— 


168 pipiens congo .... 


— 


— 


— 


— 


16 


Summer 




— 


— 


170 bulliens ..... 


— 


— 


— 


— 


— 


— 


— 


— 


— 


171 emini bailundensis . 


— 


— 


— 


— 


— 


— 


— 


— 


— 


172 lais namba .... 














_ 


_ 




_ 


_ 


173 rufilata ansorgei 


— 


— 


— 


— 


— 


— 


— 


16 


— 


174 textrix mystica 


— 


— 


— 


— 


— 


— 


i<J 




— 


bulubulu 


— 


— 


— 


— 


— 







i6,i9 





175 aridula lobito .... 


IV 


— 


— 


— 


— 













176 dambo ..... 


— 


— 


— 


2 6*. 1 $ 
(2) 


3? (1) 


Summer 


— 


— 


— 


180 Melocichla mentalis grandis . 














_ 


_ 


_ 


_ 


182 Hirundo albigularis ambigua . 


— 


— 


— 





— 


— 


16, i 9 


— 


— 


183 dimidiata marwitzi . 


— 


— 


— 





— 


— 




— 





184 senegalensis monteiri 


— 


— 


— 





— 


— 


io* 


— 


— 


185 s. semirufa .... 


— 


— 


— 





— 







i9 





186 nigrorufa .... 


— • 


— 


— 





1 6, 2 ? 


Summer 


— 


16 


— 


187 angolensis .... 


— 


— 


— 







— 


— 




— 


188 griseopyga .... 


— 


— 


— 


M 


— 


— 


— 


— 


— 


189 Riparia c. cincta 


— 


— 


— 



















190 Ptynoprogne fuligula rufigula 


— 


* 


— 





— 











— 


191 Petrochelidon rufigula .... 


— 


* 


— 





— 


— 


3 6, 6 9, 


— 


— 


202 Lanius collaris capelli .... 





_ 


_ 


I«J 






(1) eggs 






257 Ploceus temporalis .... 


— 


* 


— 










1 6, 1 







262 Euplectes orix orix (?) . 


— 


— 


— 


2? 


— 


? June 







z 


263 capensis zambesiensis 


— 


— 


1 




— 


? June 










angolensis 




























265 Coliuspasser axillaris bocagei . 


— 


— 


— 


— 


2 6\ 1 9, 
2 imm. 


? May /June 


— 


— 


— 


266 h. hartlaubi 


— 


— 


— 











1? 





— 


267 a. ardens .... 


— 


— 


1 imm. 6 


— 


— 


June 




— 


— 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 387 

Reeds and Open Country 



Districts 




Mt 


. Moco 




South-western Districts 


Western Districts 


7 


9 


20 


Breeding 


8 


Breeding 


10 


11 


13 


Breeding 


15 


17 


18 


Breeding 


- 


- 


- 


J^y 


- 


- 


— 


- 


- 


- 


- 


1 juv. $ 

I? 


- 


Aug. 
? Oct. 

July 


5 

7 
8 

10 
12 





- 





- 


- 








io\ 1? 


i<J 


?Oct. 


= 


- 


18 
26 
38 
39 
42 
64 

























i<J 


?Oct. 





iimm.6" 









i<? 


- 


- 


?Oct. 


- 


- 


= 


- 


- 


- 


- 


- 


- 


- 


65 


- 


1? 


- 


Summer 


- 


- 


- 


= 


- 


- 


- 


- 


- 


- 


66 
69 
70 



































i<? 








72 


1 <J, i? 























— 


_ 





— 


— 





73 
74 




— 


— 


Summer 


— 


— 


— 


— 


1 6*. 2? 


?Oct. 


— 


— 


— 


— 


i<J 


— 


— 


?Oct. 


— 


— 


— 


— 




— 


— 


— 


— 





75 




— 


— 


? Oct. 


— 


— 


— 


— 


— 


— 


— 


— 


— 


— 


78 


- 


x<J 


- 


?Oct. 

Summer 

? July 


- 


- 


- 


- 


- 


- 


- 


= 


- 


E 


79 
80 
81 


- 


— 


— 


? Sept. 
? Sept. 


1? 


? Sept. 


1 <J 


— 


— 


— 


- 


- 


— 


- 


106 
122 
126 
127 











Aug. 




















i? 








?July 


128 
141 


























r? 


?July 














145 


— 


— 


— 


— 


— 


— 


— 


— 


— 


— 


— 


— 


— 


— 


148 


1? 


— 


— 


Summer 


— 


— 


z 


— 


z 


z 


z 


— 


— 


— 


160 
161 


1 (J, 2? 


z 


z 


Summer 


z 


z 


— 


— 

















z 


162 


— 


— 


— 


Summer 


— 


— 


— 


— 


— 


— 


— 


— 


— 


— 


164 


— 


— 


— 


Summer 


— 


— 


— 


— 


— 


— 


— 


— 


— 


— 


165 


2 (J, 1 


— 


I<? 


Summer 


I ?, 2 


Summer 


— 


— 


— 


— 


— 


— 


— 


— 


167 


— 


— 


— 


— 


— 


— 


— 


— 


— 


— 


I0*,2? 


1 imm. 


16* 


— 1UO 

July onwards 170 




— 


— 




2?, I 


Summer 








- 


~ 


"~ 


"" 


— 


171 


K?(I) 


— 


— 


Summer 


3? 


Summer 


— 


— 


— 


— 





— 


— 


— 


172 


— 


— 


— 


Summer 


— 


— 


I<? 


— 


— 


Summer 





— 


— 


— 


173 





— 


— 


Summer 
Summer 

? Sept. 
? Sept. 
? June 
? Sept. 
Summer 


— 


— 


— 


— 


— 


— 





— 


— 


— 


174 


I<? 


- 


- 


- 


- 


- 


- 


- 


- 


- 


16" 


- 


Summer 


175 
176 

180 
182 


— 


I? 


— 


- 


— 


- 


- 


— 


— 


— 


— 


— 


- 


183 
184 
185 
186 


I? 


— 


— 


? Sept. 


— 


— 


— 






- 


- 


= 


- 




187 
188 
189 
190 
191 


*<?, 


— 


— 


Aug. 


1 


— 


— 


— 


— 


— 


— 


— 


— 


— 


2$, I 




























202 








? Sept. 








— 


— 


— 


— 





_ 


— 


_ 


257 


— 


— 


— 


— 


— 


— 


— 


— 


— 


— 


— 


— 


— 


— 


262 
263 

265 


— 


16* 


- 


?Ju"iy 


1 


?July 


— 


— 


— 


— 


— 


— 


— 


— 


r? 


- 


— 


?July 


— 


— 


- 


- 


- 


- 





- 


- 


- 


266 
267 



388 THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 

Table III. — 





Other 
references 


Eastern Districts 


Central 




Tables 


Syst. 
Notes 


1 


2 


3 


Breeding 


4 


5 


6 


268 Coliuspasser progne ansorgei . 

272 Ortygospiza atricollis minuscula 

275 Pytilia afra ...... 

277 Lagonosticta rubricata congica 

278 landanae .... 

279 jamesoni ansorgei 

280 nitidula .... 

281 Clytospiza c. cinereovinacea . 

282 Coccopygia melanotis bocagei 

283 Estrilda astrild niediecki 

angolensis 

284 paludicola benguellensis 
286 Uraeginthus bengalus katangae 
290 Serinus mozambicus tando 

292 flavivertex huillensis . 

293 atrogularis lwenarutn . 


- 


* 


1?, 
1 imm. 

1? 


1 <J, 1 ?, 

1 imm. 0* 
2 6" (1) 


1 3 


? May /June 
? June 
June 

? Sept. 


1? 


16* 

2? 


1? 



Table IV. — Birds 


of the Thornbush, Semi-desert, 




Other references 


Thorn, 






Syst. 








Tables 


Notes 


11 


12 


3 Francolinus sephaena zambesiae .... 




* 






8 Pternistes a. afer 


III 


— 


— 


— 


9 Numida mitrata (head only) .... 


— 


— 


— 





n Rhinoptilus africanus bisignatus .... 


— 


— 


— 


— 


13 Eremialector bicinctus ansorgei .... 


— 


— 


— 


— 


20 Agapornis roseicollis catumbellae .... 


— 


— 


— 


— 


23 Corythaixoides concolor pallidiceps 


— 


— 


2<J 


— 


32 Tockus monteiri ....... 


— 


— 


— 


— 


33 Otus scops senegalensis 


— 


* 


— 


— 


35 Bubo a. africanus 


— 


— 


— 


I<? 


46 Colius indicus lacteifrons 


— 


— 


— 




47 castanotus ...... 


1,11 


— 


— 


— 


51 Tricholaema 1. leucomelas ..... 


— 


— 


— 


— 


54 Pogonolius chrysoconus extoni .... 


I 


— 


— 


— 


55 Indicator minor damarensis 


II 


* 


— 


— 


58 Campethera abingoni annectens .... 


— 


— 


— 


— 


61 Dendropicos fuscescens hartlaubii ^ stresemanni 


I 


* 


Id\ I? 


— 


67 Mirafra passerina ....... 


— 


— 


— 


— 


08 sabota ansorgei 


— 


— 


— 


— 


71 Spizocorys starkii 


— 


— 


— 


— 


72 Eremopterix verticalis damarensis 


Ill 


— 


— 


— 


81 Turdoides leucopygia hartlaubii .... 


III 


— 


I? 


— 


85 Pycnonotus nigricans 


— 


— 


— 


— 


96 Alseonax c. cinereus ...... 


— 


* 


— 


— 


100 Parisoma subcaeruleum ansorgei .... 


— 


— 


— 


— 


101 Bradornis pallidus murinus ..... 


I 


— 


M 


— 


102 infuscata benguellensis .... 


— 


— 


— 


— 


108 Batis pririt ....... 


— 


* 


— 


— 


120 Monticola brevipes kaokoensis .... 


— 


— 


— 


— 


123 Oenanthe monticola albipileata .... 


— 


— 


— 


— 


124 Cercomela familiaris angolensis .... 


I 


— 


— 


— 


129 Karrucincla schlegelii benguellensis 


— 


— 


— 


— 


130 Xenocopsychus ansorgei 


— 


* 


— 


1 6 


138 Erythropygia leucophrys munda .... 


— 


— 


I? 


— 


ovamboensis . 


— 


— 


— 


— 


140 paena benguellensis .... 


— 


— 


— 


— 


144 Achaetops pycnopygius ..... 


— 


— 


— 


— 


146 Calamonastes fasciolatus pallidior .... 


— 


— 


— 


— 


151 Apalis f. flavida 


— 


— 


I? 


— 


154 Sylvietta rufescens ansorgei ..... 


— 


* 


— 


— 


mossamedes .... 


— 


— 


— 


— 


155 Eremomela icteropygialis puellula .... 


I 


* 


— 


— 


159 Camaroptera brevjcaudata sharpei (?) 


I, II 




(2) 





THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 



389 



Continued 



Districts 


Mt 


Moco 


South-western Districts 


Western Districts 


7 


9 


20 


Breeding 


8 


Breeding 


10 


11 


13 


Breeding 


15 


17 


18 


Breeding 


- 


- 


- 


- 


- 


- 


- 


- 


- 


- 


- 


- 


- 


— 268 


16* 


- 


- 


?Ju~iy 


- 


- 


- 


- 


- 


- 


- 


= 


- 


— 272 

— 275 

— 277 






































I(J 


?July 278 


— 


1 


— 


?July 
June 


— 


— 


— 


— 


1? 


?July 


— 


— 


— 


— 279 

— 280 

— 281 


1 c?, 1 9, 


— 


2d\ 


June 


— 


— 


— 


— 


— 


_ 


_ 


2 imm. 








2 imm. 




















2^,1? 


— 


— 


June 


2? 


? June 


— 


— 


1 c? 


? June 


— 


— 


— 


— 282 


2 imm. 








? June 





























— 283 


1 a, i? 


— 


— 


? June 


— 


— 


— 


— 


— 


— 


— 


- 


- 


— 284 

— 286 

— 290 


23, i? 


— 


— 


? June 


— 


— 


— 


— 


_ 


— 


— 


_ 


_ 


— 


— 


— 


— 


1? 


? Sept. 


— 


— 


— 


— 


— 


— 


— 


— 292 




























— 293 



Tangled 


Thickets and Rocks of 


Western Districts 








Thickets and Rocks 


Semidesert 


13 


15 


17 


19 


Breeding 


14 


16 


Breeding 


















&i B 


1? 


— 


— 


— 


Summer 


— 


— 


— 


3 


1 <J, a 9, 


— 


— 


— 


July/Aug. 


— 


— 


— 


T:8 


1 juv. $ 
















El 


— 


— 


— 


— 


— 


1 imm. 


— 


— 


9 


— 


— 


— 


— 


— 


— 


Ic? 


?July 


11 


— 


— 


— 


— 


— 


ic? 


— 


?Oct. 


13 


1$, 2 imm. <? 


— 


— 


— 


— 


— 


— 


— 


20 


1 imm. $ (1) 


















— 


— 


— 


— 


Summer 


— 


— 


?July 


23 


— 


— 


i<J 


— 


Summer 


— 


— 


32 
33 


— 


— 


— 


— 


Summer 


— 


— 


— 


35 


i<? 


— 


— 


— 


?July 


— 


— 


— 


46 


— 


— 


ic? 


— 


?July 


— 


— 


— 


47 


— 


— 




— 




— 


I? 


Summer 


5i 


i<? 


— 


— 


— 


?Oct. 


— 


— 


— 


54 


1? 


— 


— 


— 


?July 


— 


— 


— 


55 


— 


— 


ic? 


— 


Summer 


— 


— 


— 


58 


— 


— 


— 


— 


?Oct. 


— 


— 


— 


61 


— 


— 


— 


— 


— 


ic? 


— 


?July 


67 


— ■ 


— 


— 


— 


— 


— 


I C?, 2 9, I 


?July 


68 


— 


— 


— 


— 


— 


— 


2 9, 1 

Ic? 


?July 


7i 
72 
81 


1 


_ 


— 


— 


Summer 


— 





2$ 


— 


— 


— 


?July 


— 


— 


— 


85 


— 


— 


— 


1 imm. $ 


July 


— 


— 


— 


96 


— 


— 


— 


— 


— 


— 


2c? 


? Aug. 


100 


I<J 


— 


— 


— 


?July 


— 





— 


IOI 


— 


— 


— 


— 




— 


Ic?, I 


?July 


102 


— 


— 


— 


— 


— 


ic? 


Ic?, 2? 


?July 


108 


2 6*. 2$ 1 imm. 


— 


— 


— 


? May/June 


Ic? 


Ic? 
Ic? 


?Oct. 


120 
123 

124 


— 


— 


— 


— 


— 


I? 





— 


— 


— 


— 


— 


— 


I? 


?July 


129 


x <J, 1 9 , 1 









?Oct. 
?Oct. 





— 


— 


130 

138 


— 


— 


1$ 








I? 


Ic? 


? July 


140 


2c? 


— 


— 


— 


?Oct. 


— 


— 


— 


144 


— 


— 


— 


— 


— 


— 


Ic? 


?July 


146 


2? 


1 imm. 


— 


— 


July 


— 


— 




151 


— 


— 


— 


— 




— 


Ic?, I? 


Summer 


154 


— 


— 


— 


— 


— 


I? 


I c? 


Summer 


155 
















~ 


159 



39o 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 



Table IV. — 









Other references 


Thorn, 






Syst. 








Tables 


Notes 


11 


12 


162 Cisticola chiniana nr. frater 


I 


* 






166 fulvicapilla hallae . 






I 


— 


— 


i? 


175 aridula lobito 






III 


— 


— 




177 Prinia subflava ovampensis . 






I 


— 


1? 


— 


179 flavicans ansorgei 






I 


— 


— 


— 


199 Prionops gabela ..... 






II 


* 


— 





206 Laniarius bicolor guttatus . 






— 


* 


— 


— 


207 atrococcineus 












1 6\ 1 9 


_ 


208 Dryoscopus cubla hamatus . 






1,11 


— 


(1) 


— 


210 Lanioturdus torquatus 






— 


— 


26*, 2? 


— 


212 Tchagra australis nr. damarensis . 






— 


— 




1? 


220 Parus afer benguelae .... 






— 


— 


— 




221 niger carpi .... 






I 


— 


— 


— 


227 Lamprotornis n. nitens 






— 


— 


— 


i<J 


228 mevesi mevesi 






I 


* 


— 


1 <? 


230 Onychognathus nabouroup benguellensis 






— 


— 


— 




241 Cinnyris fuscus ..... 






— 


— 


— 


— 


250 Plocepasser mahali ansorgei . 






— 


— 








251 Passer motitensis benguellensis 






— 


— 


— 


— 


253 diffusus georgicus 






— 


— 


— 


— 


255 Sporopipes squamifrons 






— 


— 


— 


— 


256 Ploceus cucullatus collaris 






— 


— 


— 


— 


264 Euplectes aurea 






— 


— 


— 


— 


274 Pytilia m. melba .... 






I 


— 


— 


— 


276 Lagonosticta senegala pallidicrissa 






— 


— 


— 


— 


287 Granatina granatina .... 






— 


— 


— 


— 


294 Poliospiza albogularis crocopygia . 






— 


— 


— 




298 Enberiza f. flaviventris 






I 


— 


i? 


~ 


299 Fringillaria capensis nebularum 






— 


— 






300 t. tahapisi 






— 


— 


— 


~ 


301 i. impetuana 


— 


— 


— 


- 



DISTRIBUTION THROUGHOUT AFRICA OF 
SPECIES FOUND IN ANGOLA 

Woodland Birds (Table I) 

The majority of birds collected in the Brachystegia woodlands belong to species, 
and even subspecies, with a wide distribution in tropical Africa. In some cases, 
as with the Groundscraper Thrush, Geocichla litsipsirupa, this extends from Abys- 
sinia to South Africa ; in others, such as the warblers Eremomela salvadorii and 
E. atricollis, it is limited to the woodland areas of higher rainfall in Angola, south- 
western Belgian Congo and western Northern Rhodesia. Other species are limited 
to the drier woodland areas of southern Africa ; these include the shrike Lanius 
souzae, and the Angola Rock Thrush, Monticola angolensis. 

Only three species listed exclusively in Table I have a more specialized distribution 
and all are highland birds. They are Francolinus finschi and the two sunbirds 
Nectarinia bocagei and N. kilimensis ; the francolin with a sporadic distribution 
in the highlands of Angola north to Brazzaville ; N. bocagei confined to the central 
highlands of Angola*, and N. kilimensis to the highlands of Angola and East Africa. 
There are also several well-differentiated subspecies endemic to the highlands of 
central Angola, the most distinct being the Angola race of the Brubru Shrike, 
Nilaus afer affinis, and of the seedeater, Poliospiza gularis benguellensis. 

* Also at Kalandji, Kwango dist., Belgian Congo. (Schouteden, Rev. Zool. Bot. Afr. 59, 
1959:326). 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 



39i 



Continued 



Thickets and Rocks 


Semidesert 


13 


15 


17 


19 


Breeding 


14 


16 


Breeding 


1 6" 








Summer 








162 


1? 


— 


— 


— 


Summer 


— 


— 


— 


166 


— 


— 


— 


— 


— 


— 


1? 


Summer 


175 


1? 


— 


— 


— 


?July 


— 


— 


— 


177 


— 


— 


— 


— 




16", 1? 


16", 1 


?July 


179 


— 


— 


— 


1? 


?July 


— 


— 




199 


1 (J, 2 9, 


1 imm. 


16" 


— 


July/Aug. 


— 


— 


— 


206 


1 imm. <J 


















1? 











?July 











207 

208 




— 


— 


— 


? Oct. onwards 


— 


— 


— 


210 


— 


— 


— 


— 


Summer 


— 


— 


— 


212 


— 


— 


— 


— 


— 


— 


16", 1? 


?July 


220 


16", 1? 


— 


1? 


— 


? July onwards 


— 






221 


— 


— 


— 


— 


?July 


1? 


— 


— 


227 


— 


— 


— 


— 


Summer 


16" 


— 


Summer 


228 


— 


— 


1? 


— 


Summer 




— 


— 


230 


— 


— 




— 


— 


— 


3 6", 2 $, 


? Oct. onwards 241 














1 imm. 






— 


— 


— 


— 


— 


1? 


— 


?July 


250 


— 


— 


— 


— 


— 


— 


16" 


?July 


251 


1? 


— 


— 


— 


?July 


— 






253 


— 


— 


— 


— 


— 


— 


1 juv. 


Aug. 


255 


— 


— 


16,1? 


— 


?Oct. 


— 


— 


— 


256 


— 


— 




— 


— 


— 


16" 


?July 


264 


2? 


— 


— 


— 


?July 


— 


— 




274 


i<? 


— 


— 


— 


?July 


— 


— 


— 


276 


ic? 


— 


— 


— 


?July 


— 


— 


— 


287 


— 


— 


— 


— 


— 


— 


16 


?July 


294 


— 


— 


— 


— 


Summer 


— 


— 


— 


298 


1? 


— 


— 


— 


Summer 


— 


— 


— 


299 


1 6*, 1 imm. 6*, 


— 


— 


— 


July 


16" 


— 


?July 


300 


1 imm. $ 


















~ 


~ 


~~ 


~ 


" 


" 


2 imm. 6" 


July 


301 



Eucalyptus plantations, common along the railway, have been included in this 
Table, but few birds were seen in them and the only ones noted as collected in 
them were the Paradise Flycatchers at Vouga. 



Forest Birds [Table II) 

The birds found in the eastern riverine forest have all a wide distribution in the 
lowland forests of tropical Africa. Many are found also in the riverine forests of 
northern Angola, outside the range of the expedition, but only three, the broadbill 
Smithornis capensis, the forest weaver Symplectes bicolor, and Cossypha natalensis 
were found as well at Chingoroi or Gabela, the last two represented by different 
subspecies. Two others, the greenbul Phyllastrephus fischeri and Cossypha heuglini 
were also in the montane forest on Mt. Moco. 

Of the thirty species collected in the montane forest on Mt. Moco ten are not 
confined to forests ; these include the Redbacked Coly, Colius castanotus, which 
has a limited distribution in western Angola, but the other nine have wide ranges. 
A further six species are widespread in both lowland and montane forest, Tauraco 
corythaix, Chlorocichla flavicollis, Phyllastrephus fischeri, Turdus pelios, Cossypha 
heuglini and Cisticola erythrops. Only the barbet Gymnobucco calvus is typically 
a species of the Guinea forests and is found also in lowland forest in Angola at 
Ndala Tando and Canzele. Three other species, Batis margaritae, Seicercus laurae 
and Cossypha bocagei, are found elsewhere only in patches of lowland evergreen 



392 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 



forest of the southern Congo and the north of Northern Rhodesia, and the Cossypha 
also at Leba. Two species are limited to forest in Angola, Francolinus swierstrai 
from Mombolo, Hanha (the type of Francolinus cruzi, presumed forest), and Humpata 
(Rudebeck), and the flycatcher Dioptrornis brunneus only from Pungo Andongo 
and Humpata (Rudebeck), though this species is clearly closely allied to the montane 
species of East Africa, D. fischeri and D. chocolatina. The Greater Double-collared 
Sunbird, Cinnyris afer, has a unique distribution, being found, like the Dioptrornis, 
at Leba and Pungo Andongo, but also with other races on the highlands of Ruwenzori, 
the eastern Congo, Nyasaland and the lowland forest in South Africa. A Table is 

Table of Distribution of Montane Forest Species 







to 








'to* 


i 

to 
5s 


« 


•«* 






* 

o 


8 <s 




to 






1 






1 

1 

►2 




<0 






1 


« 73 


t 

O 




o ^ 


SU 


^ 8 


Oh 


cs 




Pq 


nj®. 


ftl 


Moco and Mombolo 


X 


X 


X 




X 




X 


X 


X 


Fernando Po 


— 


X 


— 




y 


. 


— 


X 


— 


Cameroon Mt. 


— 


X 


y 




y 




y 


y 


y 


Mts. of north Cameroons 


— 


X 


y 




y 


. 


y 


y 


y 


Ruwenzori area . 


y 


y 


y 




— 




— 


— 


X 


Mts. of Kivu and L. 


y 


y 


y 




y 




— 


— 


X 


Edward 




















Mts. east of L. Tan- 


— 


— 


— 




X 




— 


— 


X 


ganyika 




















Mts. of Abyssinia 


y 


— 


— 




y 




— 


— 


— 


Mts. of East Africa 


y 


y 


— 




y 




y 


y 


y 


Mts. of S. Sudan . 


— 


— 


— 




y 




— 


— 


— 


Mts. of north Nyasaland 


y 


y 


— 




y 


. 


y 


— 


— 


Mts. of south Nyasaland 


— 


y 


— 




y 


. 


y 


— 


— 



x, same subspecies as on Mt. Moco. y, different subspecies from Mt. Moco. 

provided to show the distribution of the other seven species, all of which are confined 
to montane forest and are representatives of the relic montane fauna discussed by 
Chapin (1932 : 285) and Moreau (1952 : 888). The Angola population is subspecifically 
distinct in only four of these species in spite of the distance separating it from the 
nearest mountains in the Cameroons and Nyasaland. 

The distribution of the birds found in the escarpment forest at Gabela and in 
the forest at Chingoroi emphasizes their northern association. Some, such as the 
shrikes Chlorophoneus multicolor and Lanius mackinnoni and the wattle-eye Dyaph- 
orophyia concreta, range from West Africa along the northern edge of the Congo 
forest to East Africa, though with by no means a continuous distribution. The blue- 
bill Spermophaga ruficapilla follows the southern edge of the forest to East Africa, 
while the akalat Malacocincla fulvescens and the greenbul Andropadus latirostris 
embrace both ranges. Two other greenbuls Chlorocichla falkensteini and Phylla- 
strephus fulviventris, the Red-crested Turaco, T. erythrolophus, and the bush-shrike 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 393 

Telephorus viridis have limited ranges in the west from the Gulf of Guinea to their 
southern limits at Chingoroi. The Crimson-wing, Cryptospiza reichenovii, is found 
unexpectedly at Gabela rather than on Mt. Moco since it is a montane forest species 
in the Cameroons, Fernando Po, and in most of East Africa. 

The most striking feature of the distribution of the Angola forest birds is the almost 
completely different avifauna of Mt. Moco and Gabela, the only species obtained by 
ourselves and /or Heinrich in both forests being the wide-ranging Tambourine 
Dove, Turtur tympanistra, the bush-shrike Telephorus viridis, and Colitis castonotus. 
This is the more surprising in view of the small percentage of Moco birds that are 
elsewhere restricted to mountains, others being found in the northern forests of Pungo 
Andongo, Ndala Tando and Canzele where most of the Gabela species are also found. 
The two localities are only 120 miles apart and the one-time montane forests of 
Mombolo were even closer to Gabela. Since the montane forest is recognized as 
being older than the lowland forest of the Congo Basin (Moreau 1952 : 908) it suggests 
that the escarpment forests may also be of a relic type but have been more subject 
to invasion by lowland species from the north than the more isolated montane forest. 

The distribution of the birds also points to interesting affinities between the forest 
remnants in the Leba and Humpata areas and the montane forest of Mt. Moco, 
and also between the montane forest and some forest patches of north-western 
Northern Rhodesia. 



Grassland Birds [Table III) 

The distribution of the grassland birds found in Luacano and Vouga is quite 
fascinating and, to me, inexplicable. Typical species of these grasslands are the three 
longclaws, Macronyx ameliae, M.fulleborni and M. grimwoodi, the widow Coliuspasser 
progne, and the Angola Lark, Mirafra angolensis, all found at both places ; the 
Spike-heeled Lark, Certhilauda albofasciata, though collected by us only at Vouga 
as been found also in the eastern districts near Vila Luso. Thollon's Chat, Myrmeco- 
cichla tholloni, was found at Vouga only, and Francolinus albogularis at Luacano only. 
All of these, when present in the area, were found within short distances of each 
other and would therefore be expected to have the same requirements and therefore 
similar ranges throughout Africa, yet each is completely different. Among the long- 
claws the yellow M. fiilleborni is the local variant in Angola, Northern Rhodesia, 
and the southern Congo of the widespread M. croceus ; the redthroated M. ameliae 
is primarily an East African species spreading westwards through Northern Rhodesia 
to its western limit in central Angola ; the larger M. grimwoodi was known only 
from north-western Northern Rhodesia until we found it first on the road to Luacano 
and later as far west as Vouga. 

The Angola Lark is confined to the highlands of central Angola, the grasslands of 
the Zambesi /Congo watershed and north-western Northern Rhodesia, and the Marungu 
highlands of the south-eastern Belgian Congo. The Spike-heeled Lark is essentially 
a bird of the dry western semi-desert areas of South West Africa, and the pans and 
grasslands of Bechuanaland and the Transvaal. It has recently been found by 
Rudebeck on the Humpata plateau and in the coastal strip north of Mocamedes. 



394 THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 

The francolin is West African, known for a long time only from scattered localities 
in the hinterland from Senegal to Nigeria, but recently discovered on the Marungu 
highlands, eastern Belgian Congo and in the Balovale area of Northern Rhodesia. 

Thollon's Chat is limited to the grasslands of the western plateaux of the French 
Congo and Angola, between the equator and 13 ° S. 

The Long-Tailed Widow has a broken distribution, being found on the highveld 
grasslands of the Transvaal and neighbouring provinces, in a strip across central 
Africa from Lake Bangweola to central and southern Angola, and in the highland 
grasslands of Kenya. 

From our own limited observations and from those of C. W. Benson (in litt.), 
and C. M. N. White (1958) in Northern Rhodesia it seems that even in very limited 
areas the grassland species are segregated to a large extent by preference for moister 
or drier conditions. Thus the Spike-heeled Lark, C. albofasciata, a species with a 
preference for sandy soil, short grass, and poor vegetation, was found at Vouga 
on the higher, and therefore well-drained, parts of the undulating grasslands east 
of the village. It may be this factor of drainage which enables these larks to tolerate 
here an unusually high rainfall, over 40 inches a year, whereas in most of their 
range the rainfall is under 10 inches a year. Similarly, lack of good drainage on some 
of the flatter grasslands of the eastern districts would account for their absence there. 

M. angolensis seems to require rather moister conditions than the Spike-heeled 
Lark, being collected at Vouga on the edges of the marshy valleys and by White 
at Balovale on the midslopes of the Minyanya plain. At Minyanya the ecological 
place of the Spike-heeled Lark on the upper slopes is filled by M. africana, which 
was found alongside M. angolensis at Luacano, where the limited amount of grass- 
land in the surrounding districts and the uniform flatness of the plains may have 
brought the species together through lack of their preferred habitats. 

The francolin, longclaws and widow-bird were associated with the dampest areas 
of the grasslands. The widows were in flocks along the ditches at Luacano, though 
in north-eastern Northern Rhodesia Benson found them associated with M. africana 
on the drier grasslands. All the longclaws were collected in similar conditions in 
very rank grass and in the marshy valleys. They seemed in Angola to be competitive 
species since all had been feeding on grasshoppers and beetles : Benson, however, 
found that the yellow species fulleborni and croceus in Rhodesia prefer drier parts 
than do the red-throated species, M. ameliae and M. grimwoodi. It may be that the 
widespread yellow species are more adaptive to differing conditions than the red- 
throated species, especially M. grimwoodi. 

Some of the francolins were found to have been eating frogs and fish, the latter 
possibly among those that had been left stranded with the drying up of the summer 
floods. It is most improbable that fish or frogs form an essential part of the diet 
of F. albogularis (others collected by Serle in Nigeria had been feeding on cassava), 
but high rainfall does seem to be a primary requirement for the species since the 
range lies entirely in the area between the 40-inch and 60-inch isohyets. 

It is easy to see therefore, why all the species of the wetter parts of the grassland 
should be absent from the dry grasslands of south-western Africa, but not apparent 
why each should have such differing ranges within the wetter areas, the broken 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 395 

distribution of Coliuspasser progne being particularly surprising. 

From our observations of Thollon's Chat it would seem unlikely to be found except 
where there are occasional small trees or bushes on which to perch, these perches 
being used as observation posts to which the birds returned time and again after 
foraging. Lack of suitable perches would account for their absence on the Luacano 
plains, and if all the grasslands of the eastern districts are similar to those we crossed, 
this may be the limiting factor in distribution of the species eastwards. 

Among other families classed in Table III are many cisticolas and members of the 
Finch family that are birds of grasses and reeds rather than open grassland. Most 
of these are wide-spread but many of them have local and very distinctive races in 
central and western Angola. 

Birds of the Western Thornbush, Semi-desert, Rocks and Tangled Thickets {Table IV) 

In contrast to the complexities of distribution among the grassland birds of the 
central and western districts the ranges of all western species follow regular and 
comprehensible lines governed by the topography of south-western Africa and the 
limits of the thornveld. Thus, with the exception of Euplectes aurea which was 
possibly introduced from Sao Tome, all the species collected in the semi-desert 
areas are found in similar areas in South West Africa, though in many cases are 
racially distinct. 

Typical thornveld species such as the Pied Barbet, Tricholaema leucomelas, and 
the gregarious larks such as Eremopterix verticalis, have a wider distribution through 
the Kalahari and have been found, or can be expected, in Angola as far north as 
Luanda (see Serle, 1955), while species more typical of the Vornamib scrub such as 
Schlegel's Chat, Karrucincla schlegeli, are unlikely to be found north of Lobito. 

Similarly, among the rocks of the Chela escarpment are such species as the Pale- 
winged Starling, Onychognathus nabouroup, |the rock-jumper, Achaetops pycno- 
pygius, and the Chat-flycatcher, Lanioturdus torquatus, all of which have limited 
ranges in the highlands of south-western Africa. The range of the indigenous 
Xenocopsychus ansorgei is even more limited, being apparently restricted to the 
rocks of the Chela escarpment and in the neighbourhood of Mt. Moco. 

The overall pattern is of the predominantly South West African avifauna petering 
out northwards along the coast and the edge of the escarpment. 

AVIFAUNAL ZONES AND SPECIATION 

Avifaunal Zones 

The distribution of birds found in the different types of vegetation has been 
discussed at some length because it seems that an understanding of this factor is 
primarily important in assessing the relationship between many forms found in 
Angola. Repeatedly it will be found that birds of many different families have 
similar specific or sub-specific ranges which lie along the same boundaries and coin- 
cide with the limits of one or another type of vegetation. 

Thus western Angola can be divided into three avifaunal zones (see Map) : — 
Zone 1. Above the escarpment and coinciding with the limits of Brachystegia 
woodland from the Sa da Bandeira area north at least to Malanje. This I shall call 



396 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 



the Brachystegia Zone and in it East and Central African species and subspecies 
predominate. 

Zone 2. The Acacia Zone, including the desert and semi-desert and poorer parts of 
the coastal plain, and the thorn country on top of the escarpment south and west of the 
Brachystegia Zone. In this South West African species and subspecies predominate. 

AVIFAUNAL ZONES OF WESTERN ANGOLA 



Cam a bo he la 



> -. / 



Ben 



Mocamedes 




) 

c 



*^ 



^ __.> 



Miles 



Zone 3. The north-western coastal lowlands where a richer and more varied 
vegetation is found than in Zone 2, and the escarpment area including the escarp- 
ment forests. This I shall call the Escarpment Zone. Little collecting was done in 
it except in the forests at Chingoroi and Gabela and therefore the distribution of 
birds, other than forest species, has not been discussed ; it would seem however 
to contain a number of interesting endemic forms about which much remains to 
be learnt. These include the two newly-discovered species Prionops gabela and 
Muscicapa gabela, both found originally in secondary forest and the Prionops since 
found in thickets below the escarpment : also the warbler, Macrosphenus pulitzeri, 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 397 

known only from the type specimen, and the forest greenbuls, Chlorocichla falken- 
steini and Phyllastrephus fulviventris. Among endemic forms are three which have 
been discussed in the Systematic Notes, Malaconotus hypopyrrhus blanchoti, Camar- 
optera brevicaudata harterti and Tchitrea melampyra bannermani. 

The Escarpment Zone may even be found to extend in pockets as far south as 
Vila Arriaga and Capangombe as has been suggested in the Systematic Notes in 
the discussion on the distribution of the two races of Lamprocolius mevesii. 

The interest of these zones lies in the fact that the zonal limitations are applicable 
to some birds that are not primarily associated with the typical vegetation ; for 
example Cercomela familiaris is a bird of rocks and kopjes rather than woodland 
or thornbush, but the two races falkensteini and angolensis are confined respectively 
to the Brachystegia and Acacia Zones. Similarly, the Black Tit, Parus leucomelas 
insignis, is found only outside woodlands in clearings or thickets but its true range 
is limited to Brachystegia Zone with an atypical population in the Escarpment Zone 
but not probably in the forest. It should, in fact, be emphasized that the zonal 
limits of the Escarpment Zone probably apply less to typical forest birds than to 
others, for several forest species are found in patches of riverine forest and in montane 
forest, within the Brachystegia Zone. 

On the Map the zig-zag line dividing Zones 1 and 2 is intended to indicate that 
the two zones interlock where patches of Acacia and Brachystegia alternate. Some 
sub-divisions of Zones 1 and 2 have been shown, which were indicated in the discus- 
sion on distribution. No attempt has been made to define the zonal limits in the 
northern and south-eastern parts of the country from which little or no information 
on the birds is available. 

Speciation 

The rapid changes of vegetation and of altitude in the west of the country which 
are responsible for forming the zonal boundaries and the pockets of forest, allow 
closely related forms associated with specialized habitats to live in proximity without 
interbreeding. These forms can either be closely allied species or subspecies. A 
typical example of representation by species is found in the Batis flycatchers, in 
which B. molitor is the representative in the Brachystegia Zone, B. pririt in the Acacia 
Zone, B. minulla in the Escarpment Zone and possibly also in riverine forest in 
the north, and B. margaritae in montane forest, while the two races of shrike, Lanius 
collaris capelli and L. c. subcoronatus provide examples of subspecies in the Brachy- 
stegia and Acacia Zones which do not apparently intergrade. 

Between good species like those of the genus Batis and subspecies like the shrikes 
there are other couplets or triplets of zonal representatives or forms ecologically 
separated whose relationship cannot confidently be defined. It would seem that 
through segregation they are in the process of developing into good species and are 
at varying stages in the process. 

In recent studies on various African birds the problem has been discussed of 
forms which behave as good species in one part of their range and as subspecies 
elsewhere, by authors such as Chapin, Moreau, Benson, White and Stuart Irwin. 
It will be found that for most of the birds considered western Angola is a critical 



398 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 



area in which speciation seems more advanced than in other parts of Africa, and it 
is hoped to develop this conception more fully in a later paper. Meanwhile the 
following list attempts to place in order some Angolan pairs or groups according 
to their degree of speciation, with the examples heading the list of forms that are 
no more than well-differentiated subspecies, grading through those of more doubtful 
relationship to some that behave as true species at the end of the list. Some of these 
pairs are zonal representatives, for the others there is some other ecological factor 
causing segregation which is indicated if possible. Reference is also made to papers 
in which the relationship has been discussed. 

The conventions of trinomial nomenclature compel one to make a dividing line 
between pairs considered subspecies and pairs considered species, although for 
forms in the centre of the list there are arguments for and against either treatment ; 
it seems impossible to attain consistency since each problem is different and is 
aggravated by lack of data on such characters as voice, behaviour, nest and eggs. 
Far more also will be known about the relationship between some of the pairs in 
the Brachystegia and Acacia Zones when adequate collecting has been done along 
their boundary in the south and south-eastern districts of Angola. 

Only in one example quoted, the Spur-fowl, Pternistes afer, does it seem possible 
that the process of speciation has been reversed and that, instead of forms evolving 
differences through isolation in different habitats, two very dissimilar forms afer 
and crainchii, which once may have appeared good species, have come together 
and hybridized and must now be considered conspecific. This suggestion of hybridiza- 
tion is put forward tentatively since this is the only case in western Angola in which 
intermediate populations are found, not apparently associated with vegetational 
changes, linking two highly differentiated forms of the thorn and Brachystegia 
country. Furthermore, these intermediate populations (to which the names benguel- 
lensis Bocage, chio Meise, and cambatelae Meise, have been given) show a degree 
of individual variation more commonly associated with hybrid populations than 
with populations of an established species. 



Examples of Stages of Speciation 



Erythropygia leucophrys manda 

(bushes in Zone i) 
Erythropygia I. ovambensis 

(bushes in Zone 2) 

Prodotiscus i. insignis . 

(Zone 3, forest ?) 
Prodotiscus i. lathburyi 

(Zone 1, highlands) 
Prodotiscus i. zambesiae 

(elsewhere) 

Zosterops senegalensis quanzae 

(highland in Zone 1) 
Zosterops s. anderssoni 

(Zone 2) 
Zosterops s. heinrichi 

(Zone 3) 



Subspecies showing intergradation. 



P. i. lathburyi a subspecific link between, 
insignis and zambesiae. 



Subspecies showing intergradation. See 
Moreau, 1957, and Systematic Notes. 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 



399 



Pternistes afer afer .... 

(Zone 2) 
Pternistes afer crainchi 

(Zone 1) 

Prinia flavicans bihe .... 

(bushes on Vouga grasslands, Zone 1) 
Prinia f. ansorgei 

(thornbush, Zone 2) 

Mirafra africana occidentalis . 

(grasslands of Zones 2 and 3) 
Mirafra a. kabalii 

(grasslands of eastern Zone 1) 

Nilaus afer affinis .... 

(highland Brachystegia, Zone 1) 
Nilaus a. brubru 

(thornbush, Zone 2) 

Laniarius luhderi amboinensis 

(escarpment forest ?, Gabela, Zone 3) 

Laniarius I. brauni 

(escarpment forest, Quicolungo, Zone 3) 

Eremomela icteropygialis polioxantha 

(woodland in Zone 1) 
Eremomela i. puellula . 

(thornbush in Zone 2) 
Eremomela salvadorii 

(woodland Zone 1) 

Lanius collaris capelli .... 

(bushes in Zones 1 and 3) 
Lanius c. subcoronatus 

(bushes in Zone 2) 

Malaconotus hypopyrrhus monteiri . 

(Zone 3) 
Malaconotus h. interpositus 

(bushes, Zone 1) 

Lamprotornis mevesii mevesii 

(mopane belt, Zone 2) 
Lamprotornis m. benguelensis 

(southern tip of Zone 3) 

Cercomela familiaris falkensteini 

(rocky hills in Zone 1) 
Cercomela f. angolensis 

(kopjes in Zone 2) 

Camaroptera brevicaudata harterti . 

(forest undergrowth, central Zone 3) 
Camaroptera b. sharpei 

(undergrowth, Zone 2 and southern Zone 3) 
Camaroptera brevicaudata near sharpei 

(undergrowth, Zone 1) 
Camaroptera b. tincta 

(undergrowth, northern Zone 2) 

zool. 6, 7. 



Well-marked forms linked through hybrid 
populations. See above, and Bowen, 1930. 



Probably well-defined subspecies, isolated 
from each other. See Irwin, 1959. 



Well-differentiated forms probably sub- 
species : widely separated geographically. 
See Systematic Notes. 

Well-differentiated forms often considered 
species but with some intergrades recorded. 



Exceptionally well-differentiated forms about 
which little is known. 



The first two well-defined subspecies ; salva- 
dorii probably a closely related species over- 
lapping in range. See Systematic Notes. 



Well-defined forms with close ranges but no 
recorded intergradation. 



Well-defined forms with close ranges but no 
recorded integradation. See Systematic 
Notes. 



Well-defined forms living in very close 
proximity ; no apparent intergradation. 
See Systematic Notes. 

Well-defined forms living in very close 
proximity; no known intergradation. 



harterti possibly a species. 
Notes. 



See Systematic 



24 



400 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 



Tchitrea melampyra bannermani 

(forest, Zone 3) 
Tchitrea viridis plumbeiceps 

Zones 1 and 2, also 3 outside forests) 

Passer griseus griseus .... 

(habitations — widespread) 
Passer diffusus diffusus 

(away from habitations, Zones 2 and 3) 

Lagonosticta landanae .... 

(Zone 3 and western Zone 1) 
Lagonosticta rubricata congica 

(eastern Zone 1) 

Parus niger carpi .... 

(Zone 3 outside desert and semi-desert) 
Parus leucomelas insignis (typical) 

(Edges of woodland, Zone 1) 
Parus leucomelas insignis (atypical) 

(Zone 3) 

Parus griseiventris .... 

(woodlands in Zone 1) 
Parus afer benguelae 

(semi-desert, Zone 2) 

Laniarius aethiopicus major . 

(undergrowth in woodlands, Zone 1) 
Laniarius bicolor guttatus and sticturus 

(undergrowth, reeds etc. in Zones 2 and 3) 

Pycnonotus nigricans .... 

(Zone 2) 
Pycnonotus barbatus 

(Zone 1) 

Elminia albicauda .... 

(woodlands and forest, Zone 1) 
Elminia longicauda loandae 

(forests, Zone 3) 

Prionops gabela ..... 

(forest and thickets, Zone 3) 
Prionops retzii 

(elsewhere) 

Turdus libonyanus verreauxi . 

(woodland and bush) 
Turdus pelios bocagei 

(forests of west) 
Turdus olivaceus stormsi 

(forests of east) 

Monticola angolensis .... 

(woodland and bushes, Zone 1) 
Monticola brevipes 

(rocky terrain, Zone 2) 



Possibly hybrid populations originally, now 
re-established as sympatric species. See 
Systematic Notes. 



Well-differentiated forms with possibly over- 
lapping ranges. See White & Moreau, 1958, 
and Benson, 1956. 

Well-differentiated forms with close ranges in 
Lower Congo : considered conspecific by 
Chapin. 

carpi a well-differentiated form living in close 
proximity to P. leucomelas. See Hall, 1959. 



griseiventris, probably good species known to 
overlap elsewhere with P. afer, but possibly 
isolated from it in Angola by the Mopane 
belt. See Hall and Traylor, 1959, and Irwin, 
1959. 

Probably good species known to overlap else- 
where through geological segregation, living 
in close proximity in Angola. See Hall, 
1954- 

Apparently good species but with possible 
hybridization in western Angola. See 

Irwin, 1958. 

Species living in close proximity on the zonal 
boundary. 



Closely allied species, but with little data on 
their relationship. 



T. libonyanus a good species ecologically segre- 
gated from the others which are possibly 
conspecific. 



Good species, closely related, representative 
of each other in their respective Zones. 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 401 

BREEDING SEASONS IN ANGOLA 

General Discussion 

From a collection such as this, taken only during the months of August and 
September and with a limited number of specimens of each species, generalizations 
on the regular breeding seasons must necessarily be limited ; nevertheless some useful 
data are provided by the presence of young birds and by the condition of the gonads 
and plumage, as to the probable breeding months of individual birds. In the Tables 
of Specimens Collected an estimate of the approximate breeding month has been 
made in all cases in which there appears to be sufficient evidence to warrant it. 
Nest with eggs, and young birds not fully grown, provide the only incontrovertible 
evidence — these young birds are listed in the Tables as " juvenile ;" young birds, 
fully grown, are classified as " immature." Some of these immature birds, still with 
considerable amounts of young plumage, also give some certainty within one or 
two months, especially when they are supported by adults in post-breeding moult. 
The estimated breeding month of these, as with juveniles and eggs, is put without 
a query in the Tables. The date of breeding of older, but still not fully adult pas- 
serines, is far more uncertain and has been indicated vaguely as ? May /June or 
? June /July to show that it probably took place at the ends of the rains (see Table 
of Rainfall) or the beginning of the dry season rather than in the early or middle 
part of the rainy season. 

Specimens in which the skull is not fully ossified are also classed as " immature ", 
but my knowledge of the length of time that full ossification takes in different 
families is too limited to allow me to make an estimate of breeding season on this 
evidence alone. Among sunbirds in particular a high proportion of apparently 
adult birds were noted as having ossification incomplete. 

The condition of gonads and plumage has also been considered in conjunction, 
though it is appreciated that both can be misleading in birds of the tropics. Those 
specimens in worn plumage with active gonads have been considered as possible 
breeders in the month or two months following collection, i.e. in the early rains ; 
those in very fresh plumage, or post-breeding moult, showing no breeding activity, 
as possible breeders two months prior to collection, and therefore during the dry 
season or winter ; those with non-active gonads, in indeterminate plumage, non- 
breeding dress, or moulting from non-breeding dress have been classified as " sum- 
mer " breeders on the assumption that they had not bred recently and are unlikely 
to do so before November. 

On these estimates it is possible to make some cautious contributions to the 
tables of breeding activity in Africa published by Moreau (1950) . It would be expected 
that Angola would fall into the regular pattern of the countries of southern Africa 
which have summer rainfall. In these countries, which include the Rhodesias, 
Nyasaland, and northern parts of the Union, the peak of breeding activity is found 
in September to December, that is in the early months of the rains and the two 
months preceding the rains, while in the dry winter months of May to August there 
is very little breeding activity except amongst birds of prey. In South West Africa, 
also an area of summer, but erratic, rains Macdonald (1957 : 30) found a progressive 



4 02 THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 

retardation of the breeding season from south to north with evidence that birds of 
Damaraland and the Kaokoveld have a peak activity from December to March. 

It was therefore unexpected to find evidence of considerable breeding activity in 
Angola in the dry winter months of June to August. Young birds of over fifty 
species were collected in August and September. These were found in all the main 
vegetational regions and were of widely assorted families. In addition nest and eggs 
were found in August of the Stone Chat, Saxicola tor quota, and the martin Petro- 
chelidon rufigula. Some rainfall figures relevant to the discussion are included in 
the Tables on p. 403 : — 

Non-passerines 

Comparatively few non-passerines were collected and these for the most part 
indicated the expected summer breeding seasons in all species in the eastern and 
central districts whether woodland, forest, or grassland, except for a juvenile 
Cursorius temminckii collected west of Munhango on 12th August, and a juvenile 
Coitus castanotus on Mt. Moco. In the western districts, in the semi-desert areas, 
the courser, hornbill and coly were in post-breeding moult : in the grasslands and 
evergreen thickets juveniles were obtained of Pternistes afer, and Francolinus leva- 
illantoides. 

The passerines are best discussed by ecological groups and districts according 
to the Tables. 

Passerines of the Savannah Woodlands {Table I) 

The flycatchers, thrushes and warblers generally indicate an August /September 
ora" summer " breeding season, sometimes slightly in advance of the corresponding 
families in Rhodesia, but similar to them in relation to the onset of the rains which 
start a little earlier in Angola. The shrikes and finch family appear irregular with 
some dry-season activity. The sunbirds, as in Northern Rhodesia and the southern 
Congo, seem frequently dry-season breeders. 

Passerines of the Forests [Table II) 

In the eastern riverine forests a number of indications of early winter or late 
summer breeding is provided by immature but nearly fully adult specimens, with 
rare indications of spring or early summer activity. On Mt. Moco, on the contrary, 
most birds had active gonads in August, and the Striped Swallow, Hirundo abyssinica, 
and Seicercus laurae were observed apparently engaged in nesting activities. The 
only winter breeder was apparently the weaver Hyphanturgus ocularis. 

In the western forests the indications were of a peak season in June /July, very 
few species collected not being represented by either a juvenile or immature specimen; 
many of the latter among the thrushes and finches were still largely in first plumage 
although the shrikes and bulbuls were apparently older, the indications of immaturity 
being in the bill colour, skull ossification, or in a few remaining immature feathers. 
It would therefore seem that in these patches of forest the birds of the trees and 
forest canopy have a breeding season slightly in advance of the birds of the under- 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 



403 



Some Rainfall Figures Relevant to the Discussion Measured in Inches, Less Than One 
Tenth of an Inch Shown as Trace " tr " 
Rainfall in 1957 (Localities on the Expedition Route) 















Year's 


Locality 


Years 


J. F. M. A. 


M. J. J. 


A. 


S. 


O. N. D. 


total 


Texeira de Sousa . 


1957 


10-2 5-9 9-6 0-3 


? — — 


? 


2-1 


4-8 3-6 8-3 


44-8 

plus 


Nova Lisboa 


1957 


6*5 5'9 6-7 2-8 


tr. — tr. 


tr. 


2-0 


2-6 9-2 13-1 


48-8 


Sa da Bandeira 


1957 


3-5 4-i 12-6 5-2 


— — — 


— 


0-4 


8-7 6-2 8-o 


48-7 


Lobito 


1957 


tr. 0-911-6 7-4 


— tr. — 


— 


0-3 


0-4 2-8 1-4 


24-8 


Gabela* 


1957 


1-5 5-3 n-8 5-5 


— — — 


— 


0-4 


2-9 4-8 5-4 


37*6 


Quimberima 


1957 


First spring rains fell 


27th September. 








Mt. Moco 




No figures available : 


first spring rains 


fell 


very locally on 24th August. 



Monthly Averages in Eastern Angola and North-western Northern Rhodesia 








Locality 


Years 


J. F. M. A. M. J. 


J- 


A. S. 


O. N. 


D. 


Yearly 
average 


Texeira de Sousa, 

3,609 feet 
Mwinilunga, 4,447 

feet 
Balovale, 3,577 feet 


1942-53 
1921-55 
1920-55 


8-8 8-6 9-3 5-2 0-5 — 
8-9 8-o 8-i 3-3 0-3 o-i 
8-5 6-9 5'8 i'2 tr. — 


tr. 

tr. 


0-3 0-7 
tr. o-6 
tr. • 3 


3*1 7'9 
3*7 7'5 
2-3 4'4 


8-4 

9-5 
8-9 


52-8 
50-0 
38-3 



Monthly Averages in Central Angola 
















Locality 


Years 


J. F. M. A. 


M. 


J- 


J- 


A. 


S. O. N. 


D. 


Yearly 
average 


Nova Lisboa, 5,577 
feet 


1940-54 


8-7 7-8 9-8 5-7 


0-4 


— 


tr. 


tr. 


o-6 5-5 9-6 


8-9 


57-o 


Ganda, 4,790 feet, 
55 m. N.E. of 


1924-52 


7-0 7-1 10-9 6-i 


5'4 


— 


— 


tr. 


i-i 6-o 9-3 


8-7 


6i-8 


Chingoroi 
Gabela,* 3,586 feet 
Quimberima, 3,700 

feet 


1947-55 
1954-57 

(Aug.) 


2-9 4-6 6-8 10-2 
4-4 i-i 7-7 8-5 


i- 4 
o-8 


— 


— 


tr. 


0-3 2-0 5-0 
1-4 5-5 io-o 


3'4 

5'8 


36-6 
45'2 



Monthly Averages in South-western Angol 


a and South West Africa 














Locality 
Coastal 


Years 


J- F. 


M. 


A. 


M. 


J- 


J- 


A. 


S. 


0. 


N. 


D. 


Yearly 
average 


Lobito 
Mossamedes 
Walvis Bay . 

Inland 
Sa da Bandeira 

5,860 feet 
Ondonga,3,593 feet 


1932-53 
I930-53 
1916-50 

1932-53 
1902-50 


o-8 1-5 
0-3 0-4 
tr. • 2 

5-9 6-2 
4-3 4*8 


4'7 
0-7 

o-3 

6-6 

3-7 


2-1 

o-5 
o-i 

3-7 
i- 4 


o-i 
tr. 
o-i 

o-i 
o-i 


tr. 
tr. 

tr. 


tr. 
tr. 

tr. 


tr. 
tr. 
o-i 

tr. 


o-i 

tr. 
tr. 

0-2 

o-i 


I -2 

tr 
tr. 

2-8 

o-5 


1 -o 
o-i 

tr. 

4'4 
i-8 


2-4 
o-i 
tr. 

6-3 
3-6 


i3'9 

2-1 

0-9 

36-2 
20-3 



* Rainfall figures for Gabela refer to the town on top of the escarpment, not to the forest on the face 
where the precipitation is higher. 

Compiled from the Meteorological Office Tables of Temperature, Relative Humidity and Precipitation 
for the World, Pt. IV, 1958, with additional figures supplied by the Meteorological Office, Harrow, and 
by Mr. O. Kroel for Quimberima. 



4 o 4 THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 

growth and bushes. Unfortunately, Heinrich (1958) has few records of breeding 
activity for the same species collected at Gabela ; such as there are conform with 
the general pattern of our collections, with young birds of Spermophaga ruficapilla 
collected at the end of July, and other species noted as having inactive gonads 
in August. On the other hand, he records breeding activity in several of the same 
species collected in similar forest further to the north at Canzele in March /April, 
most notably young birds then being found of Dyaphorophyia concreta, Lanius 
mackinnoni and Camaroptera brevicaudata. 

Passerines of Grasslands, Reeds and Open Country [Table III) 

In the east and centre the larks, chats, pipits and long-claws of the grasslands 
at Luacano, Lake Dilolo, Vouga and along the road mostly appear to be summer 
breeders. The exceptions were the Red-Capped Lark, Calendrella cinerea, of which 
juveniles were collected early in August, and Saxicola torquata which was nesting 
at Vouga. The Cisticolas, without exception, were in non-breeding dress or in moult 
to breeding dress, showing themselves summer breeders, as Lynes found. Nests 
and eggs of Petrochelidon rufigula were found on the 10th August in contrast to the 
record from the Belgian Congo of breeding in April, and a specimen of Hirundo sene- 
galensis in moult also suggests dry season breeding. The most surprising records how- 
ever concern the bishops, widows and waxbills, among which are many young birds 
and adults moulting into non-breeding dress indicating a breeding season continuing 
at least into June. The same species in the southern Congo and Northern Rhodesia 
are mostly recorded as summer breeders finishing in May at the latest. 

It is difficult to account for the contrasts and the similarities shown by different 
families in comparison with what is found in their relatives in adjacent countries, 
for example why the cisticolas should be normal and the bishops abnormal. The 
grass burning must inevitably affect the grassland birds ; this had mostly been 
completed in the east but was being carried out in August in the central highlands. 
It may be significant that it is the seed-eaters which had apparently finished breeding 
before the fires while the insect-eaters bred after the fires. 

In the western grasslands among the few specimens collected was a young Mirafra 
africana in September, as well as the young Francolin already referred to. 

On Mt. Moco the waxbills and cisticolas living in the rank grass of the mountain 
gullies also had contrasting breeding seasons, the waxbills breeding in winter (a 
number of young birds being obtained in August) and the cisticolas in summer. 

Passerines of the Western Thornbush, Thickets and Semi-desert, (Table IV) 

The number of specimens collected in the thornbush and tangled thickets on top 
of the escarpment in the south-west and on the road north from Lobito was limited, 
but it is apparent that the breeding season is very varied among the different species. 
In addition to the numbers of young spur-fowl seen and collected, young of Laniarius 
bicolor, Fringillaria tahapisi, Apalis flavida and Alseonax cinereus were collected 
in August, but other species, even in the same families, gave evidence of being 
summer breeders. 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 405 

The pattern in the semi-desert of the coastal plain is very different, with a high 
proportion of species in very fresh plumage or post-breeding moult. In arid country 
such as this the plumage fades and abrades very quickly so that when a specimen 
is found to have very fresh colours and unabraded feathers it cannot long have 
completed moult and the breeding season can therefore be assessed with greater 
assurance than with birds living in more sheltered conditions. These birds collected 
in September seem likely to have bred in June /July, and are in comparable plumage 
to those collected in the Kaokoveld in May. This three months' difference was 
unexpected, since the dry coastal belt of Angola seems a natural continuation of 
the Namib and Vornamib of South West Africa with no natural barriers between 
(the break made by the Cunene can hardly have significance). It is too long also 
to consider as a further logical step in the retardation of the breeding season from 
south to north as noted by Macdonald. 

However, the explanation may possibly lie in the rainfall figures for 1957, which 
show that in March and April there were exceptionally heavy rains at Lobito, 
1 1 -6 inches and 7*4 inches against an average over twenty- two years of 4-7 inches 
and 2-i inches for the same months. The figures at Benguela are not so high, 7-6 inches 
in March and 3-3 inches in April, but are still exceptional for that coast. It seems 
quite probable that there would be heavy casualities among nests, eggs and young 
birds under these conditions, and that second broods were therefore reared. Data 
on breeding seasons in other years on this coast is badly needed before the problem 
is really understood ; a little is provided by specimens in the British Museum collected 
by Ansorge in 1905 among which are some of Erythropygia paena, Plocepasser mahali 
and Passer motitensis in post-breeding moult at the end of July, suggesting an earlier 
breeding season in 1905 than in 1957. 

NEW RECORDS FOR ANGOLA AND EXTENSIONS OF RANGES 

Species not Hitherto recorded from Angola 

Francolinus albogularis meinertzhageni Luacano. 
Francolinus sephaena zambesiae Leba. 
Mirafra passerina Cahinde. 
Macronyx grimwoodi Luacano area, Vouga. 

Extension of Ranges in Angola 

Poicephalus meyeri matschei Luacano (western extension). 
Caprimulgus natalensis mpasa Lake Dilolo (western ext.). 
Andropadus virens zombensis Luau R., Luacano (western ext.). 
Alseonax c. cinereus 40 m. S. of Mumbondo (southern ext.). 
Cossypha h. heuglini Luacano (western ext.). 
Erythropygia leucosticta reichenowi Chingoroi (southern ext.). 

Eremomela icteropygialis polioxantha Vouga, and 100 m. West of Vila Luso (western 
ext.). 

Prinia subflava ovampensis Ouipungo, Leba (northern ext.). 
Campephaga q. quiscalina Chingoroi, (southern ext.). 



4 o6 THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 

Prionops gabela 40 m. S. of Mumbondo (second known locality). 
Chlorophoneus nigrifrons manningi Luau R., (western ext.). 
Cinnyris s. superbus Chingoroi (southern ext.). 
Cyanomitra verticalis hannermani Luacano (western ext.). 
Euplectes orix subsp? Lake Dilolo (northern ext.). 
Lagonosticta rubricate/, congica Luau R., (western ext.). 
Estrilda astrilde niediecki Vouga (northern ext.). 
Hypochaera nigerrima Chingoroi (southern ext.). 

New Races 

It was found necessary to describe three new races, mainly on specimens from 
this collection. These were published in short papers as follows : — 

Prodotiscus insignis lathburyi Hall, Bull. B.O.C. 78, 1958, p. 152 : Mt. Moco. 

Mirafra angolensis antonii Hall, loc. cit. p. 153 : 5 m. east of Luacano. 

Parus afer benguelae Hall & Traylor, Bull. B.O.C. 79, 1959, p. 12 m. SE. of Benguela. 

Sight Records 

Among species of interest seen but not collected were the Palm-nut Vulture, 
Gypohierax angolensis, common along the road between Lobito, Novo Redondo 
and the foot of the escarpment at Gabela. Hartlaub's Duck, Pteronetta hartlaubii, 
a pair on the Luau River ; the Bustard Eupodotis senegalensis mackenziei, at Vouga ; 
the Rufous-Bellied Heron, Erythrocnus rufiv enter, about twenty-five of which spent 
the day in high reeds near a river at Lake Dilolo, flighting down to the main lake 
half a mile away at sunset. A few were also seen on a small lagoon on the Humpata 
plateau near Jau. 

The Giant Blue Turaco, Corytheola cristata, was seen on the Luau River and at 
Gabela, both localities representing the southernmost limits of its range as known 
at present, the Luau River being an extension of the range shown by Moreau (1958 : 
map 4, p. 72). 

Three other species seen at the southern limits of their ranges are Monteiro's 
Twinspot, Clytospiza monteira, at Gabela, the Shining Blue Kingfisher, Alcedo 
quadribachys, on the Luau River, and Boehm's Spinetail, Neafrapus bohmi, 80 m. 
south of Luanda. 

SYSTEMATIC AND FIELD NOTES ON SELECTED SPECIES 

In working on the systematics and taxonomy of the Angola collection the standard 
works on the birds of the neighbouring countries have all been consulted, Chapin 
(1932-1954) for the Belgian Congo, Benson & White (1957) for Northern Rhodesia, 
Hoesch & Niethammer (1940) and Macdonald (1957) for South West Africa, Roberts 
(1940) for South Africa — as well as the manuscript of Mackworth-Praed & Grant's 
" Birds of the Southern Third of Africa." 

It is inevitable that all these authors do not always agree on systematic questions, 
especially the vexed one of " lumping " in genera and species. I have followed no 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 407 

author consistently, using my own preference where there are differences. On the 
generic and specific levels I am conscious that I have not always been able to study 
the problems as deeply as is necessary to form a. considered opinion ; I have perhaps 
been influenced by my mistrust of over-enthusiastic lumping, which I feel has 
sometimes been proposed to evade the necessity of making a decision. It is not, 
to me, necessarily sufficient reason for uniting two genera, that one species may 
show characters of both. 

The subspecific status of all forms listed has been studied on the evidence of speci- 
mens in the British Museum, supplemented in many cases by information and loans 
from other Museums, most notably from Mr. M. A. Traylor and the Chicago Natural 
History Museum, where a large part of the Angola collection from the American 
Museum of Natural History is temporarily housed. 

The Systematic Notes include any revisions that appear necessary of the accepted 
range of status of the various forms, and some notes on moults and plumage of the 
less well-known species. 

Most of the field-notes on sunbirds were supplied by John Williams, and the majority 
on all other birds by Tony Archer and General Lathbury. Williams was also responsi- 
ble for most of the detailed notes on food, derived from the contents of stomachs 
and crops ; but in the case of the turacos, some francolins, and Xenocopsychus, 
detailed analysis of the seeds and insects was made in the Botanical and Entomological 
Departments of the Museum. 

Since the collection was made papers have been published on collections from 
Angola by Heinrich (1958), and Meise (1958). These contain field or systematic 
notes on many of the species we obtained and in selecting species in the following 
notes I have endeavoured to avoid duplicating information and to concentrate on 
those in which some contribution can be made to present knowledge of the bird in 
the hand or in the field. Data on all other specimens are, however, readily available 
in the British Museum should they be required. 

With so many standard works now available on African birds it has not seemed 
necessary to provide other reference than the author's name and date to the forms 
discussed unless they have been recently described or the names newly brought into 
use. Type localities and measurements are included only when relevant to the 
discussion. 



2. Francolinus swierstrai (Roberts), Ann. Trans. Mus. 13, p. 72, 1929 : Mombolo, 
Cuanza Sul. (syn. Francolinus cruzi Themido 1937 : Hanha, Benguela dist.). 

Specimens 

1 $, 1 imm. cJ. Mt. Moco 7,000 feet, i7~22nd Aug. Bill and legs orange red ; 
iris hazel. Wing ad. 184, imm. moult ; bill 32, 30 ; tail 103, 84 mm. 

Field Notes 

This rare francolin was found in or near pockets of forest on the upper slopes. 
One was found pecking about among fallen leaves in the undergrowth, the other 



408 THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 

was flushed from the bank of a stream covered in thick bracken. It uttered a shrill 
harsh cry, not unlike that of Francolinus jacksoni O-Grant, and perched in a low 
tree. The crops contained seeds of two species of Leguminosae, one of which was 
a Dolichos or Neorautanenia, the other unidentified. 

Systematic Notes 

One of these specimens is an adult male with a long spur ; it is in worn plumage 
with some fresh feathers coming in on the head. The second is not quite fully adult, 
having a short blunt spur and some rusty edges to the wing-coverts, as described 
by Roberts for the type, which was also a young male. The young bird is mostly 
in fresh plumage with the primaries in moult and some fresh feathers coming in on 
the head. The gonads of both specimens were slightly enlarged. 

3. Francolinus sephaena zambesiae Praed 1920 : Mesanague, Zambesi, P.E.A. 

Specimen 

1 $ Leba, 2nd Sept. 

Systematic Notes 

A soil-stained specimen but matching well in markings with the type and others 
of this race, of which F. s. thompsoni (Roberts) is considered a synonym. 

4. Francolinus fins chi Bocage 1881 : Caconda, Huila dist. 

Specimens 

1 cJ 1 $ Mt. Moco 7,000 feet, i8-2ist Aug. 

Field Notes 

These birds were collected and others were seen, often in pairs, in the Brachystegia 
woodlands on the upper slopes of the mountain, and on the bare slopes above the 
tree line. The female was one of a pair pecking about in burnt grass and leaves. 
The male had been feeding on buprestid beetles, larvae and beans. 

Systematic Notes 

These specimens have been compared with a male and female from Brazzaville, 
lent by the Museum d'Histoire Naturelle, Paris, and with a male in the British 
Museum from the watershed of the rivers Cuanza and Luando. The three latter are 
very similar in general appearance and in the extent of rufous in the wings. The 
Moco birds in comparison have less rufous in the wings, with none on the outer web 
of the first four primaries, and the rufous on the inner web not reaching to within 
20 mm. of the tip on the first five primaries. It is possible that these differences 
may represent geographical variation. 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 409 

7. Francolinus albogularis (?) meinertzhageni White, Bull. B.O.C. 65, 1944 
p. 7 : Kumanu Plain, west Balovale, N. Rhodesia. 

Specimens 

2 c? 3 9 Plains near Luacano, 31st July-ist Aug. 

Field Notes 

Three of these White-throated Francolins were shot out of a covey of seven on 
an open grassy plain with no vegetation over a foot high. The others were a pair 
on a similar plain ; the female did not run or fly after the male had been shot. 
They had been feeding variously on beetles, grass seeds, small fishes and a small 
frog. 

Systematic Notes 

All are in worn plumage and stained with burnt grass. Even allowing for this the 
males appear to be darker, less rufous and more patterned with grey than the type 
of meinertzhageni. They show, in fact, much the same characters that distinguish 
the Luacano race of Mirafra angolensis, M. a. antonii, and it is possible that when 
better series are available from Angola and Rhodesia the Luacano francolins may 
also be found to represent a new race. 

21. Tauraco corythaix schalowi (Reichenow) 1891. 

Specimens 

2 9 Mt. Moco 7,000 feet, 16th Aug.: 1 9 Leba, 2nd Sept.: 1 $ Chingoroi, 9th Sept. 

Field Notes 

It seems worth recording that at Chingoroi, where both T. c. schalowi and the 
Red-crested Turaco, T. erythrolophus , were found together, the crop of each was 
found to contain quite different types of seeds, that of T. c. schalowi being full of 
small black seeds, probably Molvaceae and possibly a species of Sida, and some 
unidentified flattened seeds ; that of T. erythrolophus seeds, possibly of the family 
Rubiaceae. 

22. Tauraco erythrolophus (Vieillot) 1819. 

Specimen 

1 Chingoroi, 8th Sept. 

Field Notes 

This specimen was collected in not very thick forest bordering the stream. Others 
were seen later in the escarpment forest at Gabela. See under T. c. schalowi for 
crop contents. 



4 io THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 

Systematic Notes 

Moreau (1958 : 104) includes Huila in the range of T. erythrolophus : he has asked 
me to make a correction to this since the specimen in the collection at Pittsburgh 
on which this was based is actually labelled " dist. Huila " and was collected the 
same day as another labelled " Chingoroi." Chingoroi lies just within the boundary 
of the Huila district : the Turaco has not otherwise been recorded further south 
than Chingoroi, nor does there seem suitable habitat for it, and that specimen must 
therefore be presumed to have come from Chingoroi. 

33. Otus scops senegalensis (Swainson) 1837 : Gambia. 

Specimen 

1 $ 20 m. NE. of Novo Redondo, 14th Sept. Wing 125 mm. 

Systematic Notes 

This is a very small Scops Owl, very sandy in tone with fine vermiculations. 
In the British Museum it can be matched for colour with a specimen from Cape 
Town and several from West Africa. It seems very doubtful therefore if there are 
grounds for recognizing races on colour in a species with so much individual variation 
and I follow Mackworth-Praed and Grant (1952 : 650) in considering the majority 
of African Scops Owls as 0. s. senegalensis. This sandy specimen refutes the suggestion 
that birds of western Angola can be recognized as a distinct grey race, 0. s. hendersoni 
(Cassin), the type of which was obtained at sea off Novo Redondo. 

36. Ciccaba woodfordii bohndorffi (Sharpe) 1884 : Semmio, Niam-Niam country, 
Belgian Congo. 

Specimen 

1 (J Luau R. 29th July. 

Systematic Notes 

Mackworth-Praed & Grant (1958 : 333) have already noted that, on the British 
Museum series of African Woodowls, there are ample grounds for recognizing 
C. w. hohndorffi as distinct from the darker C. w. nuchalis (Sharpe) of West Africa. 
This specimen fits well with the series of bohndorffi being a bright rufous, lighter on 
the mantle than any West African bird examined. 

37. Semeiophorus vexillarius (Gould) 1838. 

Specimen 

1 J Vouga, nth Aug. 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 411 

Systematic Notes 

An unusually early record for a Pennant-winged Nightjar in the southern part 
of the continent. The pennants are still in sheath but nearly full grown (638 mm.). 

38. Caprimulgus natalensis (l)mpasa Smithers, Bull. B.O.C. 74, p. 84, 1954 : 
Mpasa, Luwingu dist. N. Rhodesia. 

Specimen 

1 (J Lake Dilolo, 4th Aug. 

Systematic Notes 

On the upper parts this White-tailed Nightjar is a very close match with one from 
Balovale and slightly less grey than others from north-eastern N. Rhodesia. Below 
it has only traces of barring on the sides of the chest, unlike the heavily barred 
Rhodesian birds, and nearer in this character to the type of C. n. fulviventris Hart- 
laub, from Bembe, which differs otherwise in being rufous with light markings above. 

39. Caprimulgus fossii Hartlaub 1857 : Gaboon. 

Specimens 

2 <J 2 $ Lake Dilolo, 5th Aug. 

Systematic Notes 

I follow Mackworth-Praed and Grant (1952 : 677) in not recognizing any races 
of the Gaboon Nightjar though I am not wholly convinced that a detailed study 
of a considerable series will support this view. From the available series it would 
appear that the population of Gaboon is consistently small with wings under 150 mm. 
and that all birds from the western parts of Africa tend to be greyer and less heavily 
marked than those from the east. At the same time individuals are found in South 
Africa as small as those of Gaboon and matching them in colour. If it is found 
necessary to recognize races our birds would probably be C. /. welwitschii Bocage. 

41. Caprimulgus poliocephalus koesteri Neumann 1931 : Sandula, S. of Cas- 
songue, Cuanza Sul/Huambo border. 

Specimen 

1 <$ Mt. Moco 7,000 feet, 20th Aug. Wing 151, bill 19, tail 113 mm. 

Field Notes 

" Hawking from a dead branch over thick grass, bush and bracken bordering 
a small stream in the late evening." The stomach contained three beetles. 



4 i2 THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 

Although the single specimen was actually collected outside the forest patches it 
has been included among forest birds in the Tables since, from the known habits of 
other races, it seems likely to be associated with the forest. 

Systematic Notes 

I believe this to be only the second known specimen of koesteri and the first known 
male. Mr. J. C. Greenway kindly compared it with the type in the Museum of 
Comparative Zoology, Cambridge, Mass. and found that it differed only in having 
the outer webs of the first and second primaries more profusely spotted with pale 
brown, in having the seven pale brown bars on the inner webs of the outermost 
tail feathers more distinct, and in having the white tip of the outer tail feather 
larger, 53 mm. against 30 mm. All these differences are consistent with sexual 
variation and he confirms my view that the type of C. koesteri, which was unsexed 
but presumed to be a male, is probably a female. 

Chapin (1939 : 413) was the first to suggest that koesteri might be conspecific 
with the Abyssinian Nightjar, C. poliocephalus Riippell, and the forms ruwenzorii 
O-Grant, and guttifer Grote, of south-west Tanganyika and Nyasaland, all of them 
being exclusively mountain birds associated with montane forest. The Moco bird 
is, in fact, a very close match with the type of ruwenzorii which differs only in having 
rather more white on the throat and tail. 

The type locality of koesteri has been given variously as " Chipepe, Bailundu," 
" Lebule, near Luimbale,'' and as " Luimbale.'' Mr. Greenway wrote me that 
Koester in a letter to Dr. Chapin in 1947 stated that the correct type locality is 
Sandula, south of Cassongue (about lat. 12 15' S.; long. 15 E.), at about 1,800 m. 

43. Heterotrogon vittatus camerunensis Reichenow 1902 : Cameroons. 

Specimens 

1 (J 1 $ Mt. Moco 7,000 feet i8-2ist Aug. Wing <J 123, $ 124 ; tail <$ 150, $ 145 mm. 

Systematic Notes 

The two races of Bar-tailed Trogon generally recognized are distinguished on 
size. These two birds, and one quoted by Chapin (1939 : 486) from Moco with a 
wing of 122 mm. are rather larger than typical camerunensis, but best placed with it. 

44. Apaloderma narina subsp. (?). 

Specimen 

1 $ 12 m. S. of Gabela, 18th Sept. Wing 133 ; tail, second innermost rectrices 
143, central pair in moult. 

Systematic Notes 

Other trogons collected at Quela, Gabela and Huila have been referred to A. n. 
brachyurum Chapin, by Heinrich (1958 : 335) but one from " Amboim " (i.e. Gabela) 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 413 

discussed by Sick (1934 : 168) is possibly intermediate between A. n. brachyurum 
and A. n. narina. Our Gabela female is short-tailed, as brachyurum, but closest to 
narina in the colour of the breast and chin, which are a clear light brown with 
rufous tinges, but it differs from most specimens of narina in having only a faint 
suggestion of the greyish band that usually separates the pink of the abdomen 
from the breast. 

45. Colius striatus congicus Reichenow 1923 : Lupungum, in Lomamu dist. 
Belgian Congo. 

Specimen 

1 <$ Luau R., 30th July. 

Systematic Notes 

This Speckled Mousebird agrees in all repects with C. s. congicus, and the colour 
of the iris was noted as being apple green. 

47. Colius castanotus Verreaux 1855. 

Specimens 

2 (J 2 $ 1 juv. <J Mt. Moco 6,000-6,500 feet : 1 c? Chingoroi 8th Sept. 1 <$ 20 m. 
NE. of Novo Redondo, 14th Sept. 

Field Notes 

The Red-backed Mousebird was fairly common on Mt. Moco in the forests and 
woodlands, it was also common where the Novo Redondo-Gabela road ran through 
hilly country with a covering of Euphorbia and thorn trees. 

Systematic Notes 

The young bird differs from the adults in having light red-brown edges to the wing- 
coverts and a dark brown, instead of a yellow-green, eye. 

48. Lybius torquatus bocagei (Sousa), Jorn. Ac. Real. Sci. Lisboa, 11, p. 958, 
1886 : Caconda, Huila dist. 

Specimens 

i(Ji? Luau R., 28-30th July : 3 <J Mt. Moco 6,000 feet, 21st Aug. Wings 4 <J 
89-94, ? 88. 

Field Notes 

One male collected on Mt. Moco was found, when collected, to have its tail bent 
sideways, presumably from incubating in a confined space since it is in post-breeding 
moult. The other two Moco birds are in worn plumage. 



4 i4 THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 

Systematic Notes 

Clancey (1956 and 1958) has reviewed the races of the Black-collared Barbet in 
southern Africa. I cannot agree with him that variation over this area is great 
enough to warrant the recognition of five races. Treating the species on broader 
lines I would recognize (1) L. t. torquatus (Dumont) from South Africa, distinguished 
by the yellow of the underparts being mixed with darker markings and vermicula- 
tions. (2) L. t. bocagei, synonym L. t. congicus (Reichenow), of Angola east to Luau 
R., with underparts less heavily marked than torquatus. (3) L. t. zombae Shelley, 
synonyms L. t. lucidiventris Clancey, and L. t. pumilio Grote, from Nyasaland, 
southern Congo, the Rhodesias, N. Bechuanaland, Portuguese East Africa and parts 
of Tanganyika, with clear yellow underparts. 

The five specimens obtained have all some dark vermiculations below and match 
well with a Caconda specimen in this respect. Above they show considerable indi- 
vidual variation particularly between the male and female from Luau R. the male 
being greyish on the mantle and coarsely vermiculated, the female brownish and 
finely vermiculated ; this variation does not seem to be sexual. 

49. (a) Buccanodon anchietae anchietae Bocage 1869 : Caconda, Huila dist. 

(b) Buccanodon anchietae rex Neumann 1908 : Duque de Braganca, Malanje 

dist. 

(c) Buccanodon anchietae katangae Vincent 1934 : Kahili R. Katanga. 

Specimens 

(a) 1 Caconda, 27th Aug.: (b) 1 <? Mt. Moco 6,000 feet, 18th Aug.: (c) 1 (J 20 m. 
N. of Lake Dilolo, 3rd Aug.: 1 (J 1 $ 28 m. N. of Vila Luso, 8th Aug. Wings (a) 95. 
(b) 1 (J 94. (c) 2 cJ 87-88, 1 ? 88 mm. 

Systematic Notes 

On these specimens the three races of Anchietas Barbet are recognizable on the 
colour and pattern of the yellow on the crown. In anchietae and rex the yellow is 
purer, less greenish than in katangae, and in rex it does not extend to the hind crown 
which is solid black. There are no topotypical specimens of rex in the British Museum 
and this is the first to answer to the description. Chapin gives the Bailundu district 
within the range of rex and it is probable that Moco is on the southern limit since 
another specimen from there in the Museum has not got the characters so well 
defined. The extent of yellow on the breast is slightly greater in katangae than in 
the other specimens. 

50. Gymnobucco calvus vernayi Boulton 1931 : Mombolo, Cuanza Sul. 

Specimens 

2 ? Mt. Moco 7,500 feet, 20th Aug. 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 415 

Field Notes 

One of these Naked-faced Barbets was collected from a party of three in a forested 
valley. It had been eating crickets and beetles but no fruit. 

53. Viridibucco coryphaea angolensis Boulton 1931 : Mombolo, Cuanza Sul. 

Specimens 

1 cJ 1 $ Mt. Moco 7,000 feet, 17th Aug. 

Field Notes 

Although this Tinker-bird is normally associated with mountain forest one bird 
was found away from the forest patches in some small isolated trees in open rocky 
country. It drew attention to itself with a soft piping call. It had been feeding on 
Loranthus fruits. 

Systematic Notes 

The female was preserved in salt : in comparison with the male it is greyer below 
lacking the yellow green wash, and above the yellow on the mantle and nape is 
paler appearing rather bleached. In my opinion these differences may be due to 
the action of the salt, since no sexual differences have been recorded in this species 
and none are apparent in a series of the nominate race. 

55. (a) Indicator minor minor % conirostris 

(b) Indicator minor damarensis (Roberts) 1928. 

Specimens 

(a) 1 (J Luacano, 3rd Aug.: (b) 1 $ Leba, 31st Aug. 

Systematic Notes 

The specimen from Luacano is interesting since it shows some approach to the 
darker more heavily streaked form conirostris, that was long considered a distinct 
species, and I have been fortunate in being able to discuss with Dr. H. Friedmann 
some of the problems of relationship between the two forms and to examine with 
him the series in the British Museum. Specimens have also been borrowed from 
Tervuren and Chicago. /. m. conirostris (Cassin), is a bird of the tropical forests 
of West Africa and the Congo and is found also in isolated pockets on Ruwenzori 
and Mt. Elgon within the range of other forms. Surrounding the range of conirostris 
are more lightly streaked forms living in savannah woodlands, /. m. riggenbachi 
Zedlitz, of West Africa to the Sudan, /. m. minor Stephens, of East and South Africa, 
and /. m. damarensis of South West Africa, the palest form. Between the two groups 
from localities encircling the range of conirostris, individuals have been taken which 
show some intermediate characters. Some of these from the base of Ruwenzori 

zool. 6, 7. 25 



4 i6 THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 

have been recorded by Chapin (1939 : 543), others from Owerri in southern Nigeria 
were described as /. conirostris pallidus Marchant. Other specimens showing inter- 
mediate characters in some degree have been examined from Tibati, N. Cameroons ; 
Ibembo, Lower Uelle ; Tembura, Bahr-el-Ghazal district, Sudan ; Kabalo, Tangan- 
yika district, eastern Belgian Congo ; Kasaji, south-western Belgian Congo ; 
Luacano ; Mt. Soque, near Luimbale, central Angola. The Luacano bird was collected 
on the edge of the riverine forest ; birds in the type series of pallidus were recorded 
as confined to high forest or broken forest ; the bird from Mt. Soque was collected 
in montane forest (Heinrich 1958 : 336) ; it therefore seems possible that all the 
intermediates are associated with some type of forest, or remnant of forest, rather 
than with woodland. This would perhaps account for the presence at Tibati of 
two apparently different forms, the intermediate specimen already referred to, 
and the type of riggenbachi. This type has not been examined since it was matched 
by Chapin (loc. cit.) with a specimen in the British Museum from the Shari River. 

There seems little to be gained by attempting to give subspecific status to the 
isolated intermediate specimens. The name /. m. pallidus could perhaps be used 
to cover all the intergrades but the Nigerian birds are closer to conirostris than 
most of the others. I prefer to leave the rest under intermediate designations 
according to their proximity, both in appearance and locality, to other races 

Other specimens examined from Angola are from Catumbela and Malanje, both 
of which are readily identified as /. m. minor. The one obtained at Leba in mixed 
scrub by the stream is, however, slightly paler and seems to match best with /. m. 
damarensis of South West Africa. It seems possible that damarensis is therefore 
associated exclusively with thorn country. 

59. Campethera cailliautii fulleborni % permista 

Specimens 

1 (J Luau R. 31st July. 1 $ near Vila Luso, 7th Aug. 

Systematic Notes 

Chapin (Ibis 1952 : 535) discusses intermediates between the barred Green-backed 
Woodpecker, Campethera permista (Reichenow), and the Little Spotted Woodpecker, 
Campethera cailliautii (Malherbe), found at Baraka on Lake Tanganyika, also 
at a locality 340 miles west of Baraka, and at Katombe (6° 29' S.: 23 58' E.). The 
Angola specimens are yet other intermediates though closer to the Little Spotted 
Woodpecker, represented in north-western Rhodesia by the race C. c. fulleborni 
(Neumann). On the mantle the round spots of typical fulleborni are replaced by 
small elongated spots : on the underparts the round black spots of the breast are 
replaced by kidney-shaped spots almost stretching to each edge of the feather, 
each one being twice as broad as it is deep. The kidney-shaped spots are more 
elongated in the male than the female. The general effect is of a bird more heavily 
marked below and more lightly marked above than fulleborni ; less heavily marked 
below and more heavily marked above than permista. The occurrence in eastern 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 417 

Angola of intergrades and ones closer to C. c. fulleborni seems logical since C. p. 
permista is found in northern Angola at Vila Salazar and C. c. fulleborni rather 
nearer, in the Balovale area of northern Rhodesia. 



60. Campethera nivosa herberti (Alexander) 1908 : Bwande, Ubangi R. Belgian 
Congo. 

Specimen 

1 <J 12 m. S. of Gabela, 18th Sept. 

Systematic Notes 

Meise (1958 : 70) has described a new race of the Buff-spotted Woodpecker, 
C. n. canzelae, from Canzele, about 200 miles north of Gabela. This male from Gabela 
is a very close match with the type of C. n. herberti ; it is the only Angola specimen 
of the species in the British Museum, but Mr. Traylor (in litt.) tells me that he, on 
specimens in Chicago, considers more northern birds to be herberti also, and that 
canzelae therefore should be regarded as a synonym of herberti. 

61. (a) Dendropicos fuscescens hartlaubii Malherbe 1849 : Zanzibar. 

(b) Dendropicos fuscescens hartlaubii $ stresemanni Grote 1922 : 
Okaukweyo, SW. Africa. 

Specimens 

(a) 1 (J Luau R., 30th July : 1 $ 20 m. SE. of Luacano, 6th Aug. : 1 ? 15 m. N. of 
Vila Flor, 25th Aug.: 1 <J 5 m. S. of Caconda, 28th Aug. (b) 1 £ 1 ? Quipungo, 
28-29th Aug. 

Field Notes 

All the specimens of {a) were collected in Brachystegia and the pair of (b) in thorn 
country. 

Systematic Notes 

The use of the name hartlaubii for widespread populations of the Cardinal Wood- 
pecker was discussed (Hall, 1956 : 101) ; these specimens together with others in 
the British Museum help to complete the picture in Angola. They show that birds 
from the Vila Salazar area in the north of Angola are consistently washed with 
yellow on the mantle, matching a series from Tanganyika ; those from the centre 
of the country are variable, the Luau R. male and a Caconda male of an older 
collection being washed with yellow although ours from Caconda is not ; those 
from the thorn country in the south show a very marked approach to the lightly 
streaked stresemanni of South West Africa this being particularly true of the Quipungo 
male although the female is more heavily streaked. 



4 i8 THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 

62. Mesopicos griseocephalus persimilis Neumann 1933 : Chipepe, Cuanza Sul. 

Specimens 

1 cJ, 2 imm. Mt. Moco 7,000 feet, 20-2ist Aug.: 1 imm. Leba, 3rd Sept. Wings 
101-106. 

Field Notes 

The Olive Woodpeckers on Moco were shot in or near the forest, one of the young 
being in a party of three. The Leba bird was one of a party of four associated with 
a mixed bird party of greenbuls and Lanioturdus torquatus, moving among the stunted 
woodland on the hillside above the gorge of the river. 

Systematic Notes 

These are the first Angola specimens in the British Museum, and on them the race 
persimilis has been accepted tentatively as distinct from ruwenzori Sharpe, since 
they are all short winged in comparison with ruwenzori. There are no colour differ- 
ences apparent between the single adult persimilis and eastern birds but it is a 
specimen which was preserved temporarily in salt and therefore not very satisfactory 
for comparison. 

64. (a) Mirafra africana kabalii White, Bull. B.O.C. 64, p. 20, 1943 : Minyanya 
Plain, Balovale. 

(b) Mirafra africana occidentalis (Hartlaub) 1857 : Gaboon. 

(c) Mirafra africana occidentalis % pallida Sharpe. 

Specimens 

(a) 2 (J 2 ? Luacano area, 31st July-ist Aug.: (b) 1 <$ 1 $ Quipungo, 29th Aug.: 
1 cJ 1 imm. <J, 60-80 m. N. of Lobito, 14th Sept.: (c) 1 $ Leba, 4th Sept. 

Field Notes 

The late Captain Grant always expressed some doubts as to whether the form 
kabalii was really a race of the Rufous-naped Lark and it was hoped to obtain some 
field evidence on this point for him. Unfortunately, at Luacano these larks were 
apparently rare and were not observed long enough for conclusions to be reached ; 
they were not heard calling. In the areas of western Angola where specimens of 
the western race were collected they were common and prominent, sitting on con- 
venient perches, sometimes uttering typical " africana " whistles, and going to 
ground and running when disturbed. The apparent difference in behaviour between 
the two races might be partly attributable to different breeding seasons, some western 
birds being in worn plumage with enlarged gonads, but must also be partly due to 
the absence of suitable perches on the Luacano plains. It seems likely that kabalii 
must have adapted itself to a more terrestrial mode of life than other forms and it 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 419 

would be interesting to observe its behaviour in the breeding season and see how it 
is modified to the rather different conditions. 

Systematic Notes 

The contrast in appearance between the specimens of the west and east of Angola 
is certainly striking. Those of the west have the long bill, rufous nape patch and 
typical lark mantle and head. The bills of the eastern birds are variable, but mostly 
short, they are almost blue-grey on the mantle with the dark centres to the feathers 
wider, to give a more scalloped effect : the rufous nape patch for the same reason 
is inconspicuous in three out of the four. 

Western birds from north of Lobito south into South West Africa show a cline 
from greyer birds in the north (occidentalis) to paler and more rufous birds in the 
south (pallida) ; the Leba specimen is intermediate, as one from the Kaokoveld 
was also found to be by Macdonald and Hall (1957 : 18) ; it seems that both represent 
steps in the cline. 

The eastern birds from Luacano match well with a topotype of kabalii. White 
(1956 : 122) refers to kabalii two specimens collected by Lynes from Missao de Luz 
(140 m. west of Luacano) and from about 190 m. further north in the Lunda district. 
They are both redder than typical kabalii and seem to me to be intermediate between 
this race and malbranti Chapin, of the French Congo, as represented by another 
Lynes' bird from Petianga, Kasai district ; malbranti in turn has close affinities with 
stresemanni Bannerman, of the Cameroon highlands and kurrae Lynes, of Darfur. 
These links between kabalii and northern races of M. africana seem to provide 
convincing evidence that it should be considered conspecific with them, all forming 
a rather distinctive group in pattern and size within the species as a whole. 

65. (a) Mirafra angolensis angolensis Bocage 1880 : Caconda, Huila dist. 
(b) Mirafra angolensis antonii Hall 1958 : 5 m. east of Luacano, Moxico. 

Specimens 

(a) 3 (J 1 $ Vouga, nth Aug.: 1 ? 50 m. E. of Vila General Machado, 9th Aug.: 
1 cJ Mt. Moco 6,000 feet 16th Aug.: (b) 6 <J 3 ? Luacano area, i~3rd Aug. 

Field Notes 

All were collected on open plains, usually near the wetter valleys or ditches 
rather than on the dry open spaces (see p. 394). They were in pairs or single. The 
flight was noted as being low and straight and birds put up flew some distance 
before settling again. One was seen to plane down from a height and perch on an 
anthill. 

Systematic Notes 

The systematics of the Angola lark were discussed when M. a. antonii was des- 
cribed. 



420 THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 

74. Anthus vaalensis neumanni Meinertzhagen 1920 : Ambaca, Cuanza Norte. 

Specimens 

1 $ Lake Dilolo, 6th Aug.: 1 $ 43 m. W. of Texeira de Sousa, 31st July: 1 $ 14 m. 
W. of Munhango, 9th Aug.: 2(Ji? Vouga, 10-nth Aug.: 1 $ Nova Lisboa, 13th 
Aug.: 1 (J Leba, 2nd Sept.: 2 $ Humpata-Jau road, 4th Sept. Wing 3 <J 92-100, 
7 ? 92-102 : hind claw <J°. 9-12 mm. Bill colour, dark brown or blackish above, 
pale below with yellow gape. 

Field Notes 

All these pipits were found in fairly open ground, often in the vicinity of water 
or marsh. On the Vouga plains they were alongside Mirafra angolensis on the slopes 
above the marshy valleys, rather than with the Longclaws in the bottom of the 
valleys. 

Systematic Notes 

The identification of the Plain-backed Pipits of Angola was discussed in a recent 
paper (Hall, 1959) in which it was shown that it is not practical to recognize more 
than one race of A . vaalensis in Angola, although there is some geographical variation. 
Of these specimens all from Vouga and further east are rather greyer than those 
from the west, the differences not entirely due to the fact that the eastern birds 
are slightly stained from burnt ground. 

75. Anthus leucophyrs bohndorffi Neumann 1906 : Kassongo, Belgian Congo. 

Specimen 

1 cJ Mt. Moco 6,500 feet, 18th Aug. Wing 1 <J 102 : hind claw 12-5 mm. Bill 
colour, blackish above, bright yellow below with darker tip. 

Field Notes 

It is not possible on this single specimen to form any opinion on the ecological 
relationship between the two Plain-backed Pipits in Angola. This bird was collected 
in a grassy patch by a stream at the base of the mountain. 

Systematic Notes 

This is one of the three rather long-winged and short-clawed representatives of 
A . /. bohndorffi found in the mountains, which were discussed in the paper on identi- 
fication. (Hall, 1959). The bright yellow of the bill was very noticeable in the fresh 
skin and may possibly be a character of breeding dress. 

76. Anthus similis schoutedeni Chapin 1937 : Kwamouth, Belgian Congo. 

Specimens 

1 Mt. Moco 6,000 feet, 18th Aug.: 1 near Vila Flor, 25th Aug.: 1 <J Caconda, 
28th Aug. 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 421 

Field Notes 

All were collected in Brachystegia woodland between 5,000 feet and 6,000 feet. 
It seems that in central Africa Anthus similis is more a bird of the woods than of 
the rocks, and both the common names applied to the species — Long-billed Pipit 
in Africa or Rock Pipit in Asia — are equally unsuited to this race, schoutedeni being 
comparatively short-billed (15-17 mm.). 

77. Anthus lineiventris Sundevall 1850. 

Specimen 

1 (J Mt. Moco 7,000 feet, 21st Aug. 

Field Notes 

This bird was shot in a low tree on the edge of light woodland and the open rocky 
slopes. Chapin (1953 : 80) gives 5,000 feet as the altitudinal limit for the Striped 
Pipit. 

78. Macronyx ameliae De Tarragon 1845. 

Specimens 

1 cJ 2 $ 1 imm. $ Luacano area, 31st July-3rd Aug.: 1 <$ Lake Dilolo, 4th Aug.: 
2 (J Vouga, io-ii Aug. 

Notes on Moults 

In the British Museum there are a number of specimens of Amelia's Longclaw 
collected by Lynes, Vincent and Benson which have good data on the condition 
of the gonads. From these, and from our birds, it is evident that in M. ameliae 
(and in M. grimwoodi as well) there is an irregular and incomplete moult before the 
breeding season. This moult, as with many pipits, may be of body plumage only 
or include some secondaries and /or the central rectrices. In the males new bright 
red feathers come in on the throat, replacing orange or buff feathers. From the one 
specimen examined in full post-breeding moult (Dedza, Nyasaland, 21st April) 
I am of the opinion that the bright red throat feathers are then replaced by orange- 
pink feathers, and that therefore some of the differences in the colour of the throat 
and abdomen are due to distinctive breeding and non-breeding plumages as well 
as to wear and individual variation, and differences in immature and adult plumages 
as discussed by White (1946 : 79). 

79. Macronyx grimwoodi Benson 1955 : Chitunta Plain, Mwinilunga, N. Rhodesia. 

Specimens 

1 <J 36 m. W. of Texeira de Sousa, 31st July ; 3 $ 1 9 Vouga 10-nth Aug. 



422 THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 

Field Notes 

The first specimen was found on an open dambo near a river, and at Vouga others 
were found in the same marshy valleys as M. amelia and M. fulleborni Reichenow. 
The flight was noted as being slow with the white wing-patch conspicuous. 

The discovery of Grimwood's Longclaw as far west as Vouga is a notable extension 
to its known range. 

Systematic Notes 

The colour of the throat in all specimens is richer than in one of the original type 
series which was described as " very pale." In all the Vouga birds new feathers 
are coming in on the throat and, in the two specimens in which this moult is most 
fully advanced, the general colour is rich salmon. These specimens also have a 
distinct pink flush on the abdomen and shoulders ; one is in body moult (see notes 
on moults of M. amelia). 

From one of the Vouga birds specimens of feather lice were obtained which were 
identified by Dr. T. Clay as of the genus Ricinus. 



84. Malacocincla fulvescens dilutior White, Bull. B.O.C. 73, p. 96, 1953 : 
Ndala Tando, Cuanza Sul. 

Specimens 

3 (J 12 m. S. of Gabela, I7-I9th Sept. Wing 71-73. 

Systematic Notes 

When dilutior was described it was compared only with the nominate race. In 
appearance it seems closer to ugandae (van Someren), but has the throat paler, 
with distinct streaks and the chest bright buff rather than grey buff. It is also smaller, 
the wings of six males being 71-76 against 76-80 mm. for seven males of ugandae. 
The Gabela specimens are smaller than those from Vila Salazar (74-76). 



89. Chlorocincla f. falkensteini (Reichenow) 1874. 

Specimens 

1 ^ Chingoroi, 9th Sept. 1 $ 1 imm. P 12 m. S. of Gabela, 17th Sept. 

Systematic Notes 

The young bird was identified as immature by the skull not being fully ossified, 
by having soft edges to the gape which was yellow, and by having a light brown, 
not " crimson " or " deep brick red " iris, but the plumage is similar to the adults. 
The stomach contained large berries which were possibly coffee. 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 423 

93. Phyllastrephus fulviventris Cabanis 1876. 

Specimens 

1 (J 1 $ 1 Chingoroi, 8-ioth Sept.: 1 c? 1 ? Gabela, 18-igth Sept. 

Field Notes 

Very little is yet known of this greenbul. These specimens were collected or 
netted in forest. One was in a party of three seen hopping about in the leaves and 
low undergrowth. Two were found to have been eating insects ; this seems to be the 
usual diet of Phyllastrephus, as noted by Chapin for the species P.fischeri (Reichenow) , 
P. xavieri (Oustalet), P. icterinus (Bonaparte), P. albigularis (Junge) and P. flavo- 
striatus (Sharpe). 

96. Alseonax c. cinereus (Cassin) 1857. 

Specimen 

1 imm. $ 40 m. S. of Mumbondo, 20th Sept. Wing 69 mm. 

Systematic Notes 

This young flycatcher still has some juvenile plumage on the head and wing- 
coverts but gives the appearance of being fully grown. By its colour as well as by 
its small size it can confidently be referred to this race and not to the larger and paler 
A. c. cinereolus (Finsch & Hartlaub), which occurs in southern Angola. 

105. Hyliota australis ? slatini Sassi 1914 : Beni, eastern Belgian Congo. 

Specimens 

1 cJ 5 m. W. of Munhango, 9th Aug.: 2 <$ 1 $ 28 m. W. of Vila Luso, 8th Aug. 

Systematic Notes 

White (1957 : 35) discusses variation in this flycatcher, with particular reference 
to the amount of white in the tail. There is very little available material of any 
races and the tail pattern of the male is the chief character on which races have been 
separated. From material in the British Museum, and information given to me by 
Miss Patterson on the birds in Bulawayo it seems that H. a. australis Shelley, with 
some white on both the inner and outer webs of the outer rectrices, is confined to 
Southern Rhodesia : H. a. inornata Vincent, from Portuguese East Africa and 
southern Nyasaland, is probably separable on having rather more white on the inner 
webs : birds from Northern Rhodesia are variable, often lacking any white, and the 
population there is apparently intermediate between australis and the wholly 
black-tailed slatini of the eastern Belgian Congo. Males from the southern Belgian 
Congo and the series from eastern Angola also are wholly black-tailed. The Amani 



424 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 



race ttsambara Sclater, has a little white on the outer web but is chiefly distinguished 
by the richer colouring of the throat and breast. 

Two eastern Congo birds were borrowed from Tervuren to compare with the Angola 
series. They were found to have rather a more purplish sheen on the upper parts, 
the Angola birds being a very sooty black, and to be more washed with orange on 
the breast. These differences could well be due to wear, the Angola series being in 
worn plumage and stained with burnt grass ; all are therefore referred for the present 
to slatini. 

108. Batis pririt (Vieillot) 1818 : Lower Orange River. 

Specimens 

1 cj 18 m. W. of Cahinde, 5th Sept.: 1 (J 2 $ 12-30 m. SE. of Benguela, n-i2th 
Sept. 

Systematic Notes 

Pririt Flycatchers from Angola average smaller than those from the rest of the 
range ; but in the absence of any other distinguishing characters the difference 
does not seem great enough to warrant them being separated as a new race. 



Comparative Wing Measurements 






Angola 


• 8 (J 53-55 


8$ 52-55 


Damaraland . 


• 56" 55-6o 


7$ 54-59 


Namaqualand 


• 86* 57-59 


5$ 56-57 


Cape Prov. . 


• 36* 56-58 


3? 56-57 


Bechuanaland 


• 66* 55-58 


8$ 54-58 


O.F.S. . 


. — — 


1? 55 



109. Batis m. margaritae Boulton 1934 : Mt. Moco, Benguela. 

Specimens 

2 (J 1 $ Mt. Moco 7,000-7,500 feet, i7~2ist Aug. 

Systematic Notes 

Some doubt has been expressed as to whether B. kathleenae White, of Mwinilunga, 
is distinguishable from margaritae. Comparison of these Moco birds with kathleenae 
show that they differ in both sexes in being darker, more blue-grey on the back, 
and the blue-black of the eye-stripe and ear-coverts continues on to the hind neck 
to form an almost complete V. In addition the female kathleenae has the chestnut 
wing-coverts less rich and has a white line from the lores to the eye which is absent 
in the female margaritae. 

In a genus such as Batis where the colour and pattern of the female is the most 
striking difference between species, I am not convinced that two forms such as 
margaritae and kathleenae with almost wholly black-and-white females should be 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 425 

united, as has been done, with Batis capensis (Linnaeus), in which the female has 
chestnut underparts and a browner grey back than the male. I prefer to consider 
B. margaritae, B. capensis and B. diops of Ruwenzori as forming a superspecies. 

112. Dyaphorophyia concreta ansorgei Hartert 1905. Cabeca de Ladroes, 
Benguela. 

Specimens 

3 (?3 ?i juv. $ Chingoroi, 8-nth Sept.: 1 g 1 $ 12 m. S. of Gabela, 12th Sept.: 
(1 $ in spirit). 

Field Notes 

The Yellow-bellied Wattle-eye was most commonly netted in thick undergrowth 
of the forests. On one occasion two pairs and a party of three were seen within 
fifty yards, on another occasion one was seen with a party of Chlorocichla flaviventris. 
Several calls were noted, a sweet whistle " pih, pih, puh, puh, puh, puhh," preceded 
by a call not unlike an alarmed Yellow- vented Bulbul ; also a whistle " phee, phee, 
pheeat." The wings make a loud snap in flight. 

Systematic Notes 

The Chingoroi series are nearly topotypical : in view of the rapidity with which 
the colours fade it seems advisable to give a detailed description of the plumages 
of both sexes and the young bird. All the adults are finishing moult and show that 
both sexes have olive green backs when fresh, which wear to grey, with no metallic 
wash except on the tail. Below the males are a brilliant orange-yellow, the females 
very slightly paler on the abdomen with a rich chestnut throat and chest. The young 
male is mostly out of the nestling plumage which was described by Meise (1958 : 76). 
It is rather paler olive green on the back than the adults, has a pale yellow abdomen 
and a faint wash of orange on the chest and throat ; the wing-coverts and secondaries 
are tipped with buff ; the tail is grey with only a faint metallic tinge. The iris 
of the adults was noted as being dark brown with a silver thread round the pupil ; 
this thread was lacking in the young bird which had a greyish brown iris. 

Meise (loc. cit.) has described a new race from Canzele in which females are greyer 
than in ansorgei. The Gabela specimens show no approach to this. 

114. Tchitrea viridis plumbeiceps (Reichenow) 1898 : Malanje. 

Specimens 

1 cJ 1 $ Vouga, 12th Aug.: 1 <J 1 $ Mt. Moco 7,000 feet, 20-2ist Aug. 

Field Notes 

The pair at Vouga were among the few birds seen in Eucalyptus plantations along 
the railway. The pair from Mt. Moco were netted in thick montane forest but 



426 THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 

others were common in the Brachystegia of the foothills, and on one occasion were 
seen mobbing a Yellow-throated Sparrow. 

Systematic Notes 
See following species. 

115. Tchitrea melampyra* banner mani (Chapin) 1948 : Ngara, Cuanza Sul. 

Specimen 

1 ? 12 m. S. of Gabela, 16th Sept. 

Field Notes 

This specimen was collected in the forest. 

Systematic Notes 

It is not proposed here to go into the vexed question of the taxonomy of the Para- 
dise Flycatchers but in order to discuss the situation that prevails in Angola it 
is necessary to define the main groups under discussion. For this it is most convenient 
to follow Mackworth-Praed and Grant (1955 : 223) in considering the group comprised 
of the forms plumbeiceps of Angola and violacea Grant & Praed as separate from 
other forms of T. viridis. In these two forms the wings and mantle are cinnamon 
in colour with no white, the head is light grey with a long crest, the underparts 
are grey with a considerable amount of white on the abdomen and the under tail- 
coverts white or washed with rufous ; males have a long tail. The other group with 
which we are concerned is the generally recognized species T. melampyra (= T. 
rufocinerea of Chapin and other authors — see Grant & Mackworth-Praed 1957), 
comprising the races batesi (Chapin) of the Camaroons, melampyra Verreaux of the 
Lower Congo and extreme northern Angola (Quela and Canzele — Traylor in litt.) 
and bannermani of north western Angola, south of melampyra. T. m. melampyra 
has the wings and mantle a bright orange rufous, the head a deep metallic blue with 
a short crest, the underparts dark grey with no white on the abdomen, and has deep 
rufous under tail-coverts and a short tail. 

Chapin in his paper on hybridization among the Paradise Flycatchers (1948 : 118) 
discusses the hybrid population bannermani, derived from inter-breeding between 
the melampyra and the plumbeiceps groups. He shows bannermani to be closer to 
melampyra but showing an approach to the plumbeiceps group in having a paler 
head, longer crest and paler underparts. He indicates that bannermani has become 
sufficiently stabilized to live alongside plumbeiceps though ecologically segregated, 
bannermani being found in more wooded or forested habitat while plumbeiceps 
keeps to more open country. All localities from which bannermani has been collected 
lie in the centre of the Escarpment Zone and our single specimen was collected 
within the escarpment forest. We did not find plumbeiceps ourselves in this area 

* Since this was written the specimen in the British Museum alleged to be the type of melampyra has 
been re-examined at the request of Dr. J. P. Chapin, and found not to be the type. It is probable that the 
name melampyra will prove indeterminate : meanwhile it is preferable to use rufocinerea for this species. 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 427 

but Ansorge collected both forms at Ndala Tando, though there are unfortunately 
no data on his specimens as to the type of country in which they were obtained. 
Chapin does not cite actual evidence of the two forms breeding alongside, and, 
in fact, the majority of plumbeiceps from north-western Angola have been collected 
in the non-breeding season, June to August, when it is known to wander widely. 
There is, however, a female plumbeiceps from Dondo collected by Ansorge in Novem- 
ber to support his view and its known breeding range come, very close to the Escarp- 
ment Zone at Malanje and Moco. This seems to present an unusual case of two 
species hybridizing, possibly through some temporary breakdown of ecological 
barriers, and then at a later date the hybrid population being again divided into 
two populations, which we now recognize as bannermani and plumbeiceps, each 
of which is strongly associated both morphologically and ecologically with one of 
the parent species although showing some influence of the other parent. 

This factor has been recognized in the discussions on bannermani which is accepted 
as a race of melampyra showing an approach to plumbeiceps, but it does not seem 
to have been appreciated that it is also important in considering the status of the 
Angola population of the plumbeiceps jviolacea group. A study of all specimens of 
this group shows that there is great variability in the extent of white on the abdomen 
and in the amount of rufous in the under tail-coverts but that specimens showing 
little white and a lot of rufous are most common in Angola and the Belgian Congo 
and less common eastwards into Nyasaland and southwards into South West Africa. 
This it seems can be directly attributed to the diminishing of the melampyra influence 
away from the zone of one-time hybridization. Since plumbeiceps was described 
from Malanje I believe it should be recognized as a variable race in which the white 
of the abdomen is usually rather restricted and the under tail-coverts usually rufous, 
and that it merges imperceptibly into violacea of Nyasaland which has usually more 
white on the abdomen and in which white under tail-coverts predominate. It seems 
doubtful if T. viridis subrufa (Salomonsen 1949) from the south east Belgian Congo, 
which was described as having dark cinnamon under tail-coverts distinct from some 
Malanje specimens in which they were white or yellowish, will be found to differ 
from a larger Angola series which shows all variations. 

It is an interesting point that plumbeiceps away from the range of bannermani 
is not apparently confined to more open country since a pair were collected in the 
Moco forest. Of these the female is unusual in having elongated central rect rices 
like the male. 



118. Turdus pelios bocagei (Cabanis) 1882 : Angola. 

Specimen 

1 (J Mt. Moco 7,000 feet, 23rd Aug. Wing 132 mm. 

Systematic Notes 

I have followed Mackworth-Praed and Grant (1955 : 241) in considering T. pelios 
a distinct species from the Olive Thrush, T. olivaceus, but believe they are wrong 



428 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 



in applying the name schuetti (Cabanis) to this species. Professor Stresemann has 
examined the types of both bocagei and schuetti and emphasizes that schuetti belongs 
to the woodland species T. libonyanus : it should therefore be placed in the synonymy 
of T. 1. verreauxi and T. p. bocagei be used for the forest bird. 

The wing length of bocagei is very variable. The difference between the Moco 
bird and a series from N. Angola in the British Museum was found to be so great 
that I asked Mr. Traylor for details of specimens in Chicago. From the combined 
series it is apparent that birds from the high country inland, Moco and Quela, are 
conspicuously larger than those of the escarpment forests : when the species as a 
whole is reviewed it may be found necessary to recognize two races in western 
Angola. 

Table. — Wing Lengths of Specimens of Turdus pelios bocagei 

Locality 
(south to north) 

Moco . 
Gabela 
Dondo . 
Ndalo Tando 
Golungo Alto 
Canzele 
Quela . 

119. Geokichla litsipsirupa stierlingi Reichenow 1900 : Iringa, Tanganyika. 





6 








? 




Unsexed 


117, 


127, 


130, 


132 


119, 


121, 123, 


123 


— 


no 








III, 


119 




118 


119 








III 






— 


116, 


117. 


117. 


117 


107, 


III 




— 


116 








— 






— 


114 








109 






— 


129 








— 






— 



Specimen 

1 $ 5 m. W. of Munhango, 8th Aug. 

Systematic Notes 

Although the Angola Groundscraper Thrushes have been separated as G. I. kosteri 
(Neumann), on slight differences of size and pattern, I can see no reason to distinguish 
this specimen from G. I. stierlingi. 

127. Myrmecocichla tholloni (Oustalet) 1886. 

Specimens 

2 (J 1 ? 30 m. W. of Vouga, 12th Aug. 



Field Notes 

Thollon's Chat was first seen by the road 50 miles east of Vila General Machado 
and again on the plains near Vouga, but it proved very wary and difficult to approach. 
The three birds were finally collected near together, two having been watched for 
some while. It was found that they made use of the few small trees on the open 
plains, returning again and again to two or three selected perches after forays on 
the ground for food. The stomach contents showed this to be largely beetles and 
grasshoppers but fragments of a small lizard were also noted. 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 429 

The alarm note was a shrill peep " almost identical with the Kenya Anteating 
Chat." When the first male was shot the female returned and fluttered about in 
the vicinity. Earlier on a wounded bird was lost and almost certainly had gone to 
ground in an antbear hole. 

It was noticed that, in life, this chat looks darker than in the hand, appearing to be 
almost black with contrasting white markings. 



130. Xenocopsychus ansorgei Hartert 1907. 

Specimens 

1 cJ 1 ? $ 1 Leba, 1st Sept.: 1 <$ 3 m. SW. of Sa da Bandeira, 6th Sept. 

Field Notes 

Braun (1956 : 41) has written a full account of his observations on Xenocopsychus 
in the Sa da Bandeira area. We found it, as he did, among the rocks and trees on 
top of the escarpment (see Plate 5, a) and in the gorge of the stream. We did not 
find it as shy of sunlight as he led us to expect, the first two being collected in the 
morning of a sunny day. Two pairs at least lived close to each other on the edge 
of the escarpment, and a pair when pursued at dusk took at least temporary refuge 
in the tangled thickets behind. Both the song and the alarm note were heard : 
the double-syllabled alarm whistle by one bird was frequently answered by a single 
whistle from the other. They moved and behaved much as a Cossypha, and in 
dimensions and pattern as well have much in common with Cossypha heuglini. 

The stomach contents were found to contain fragments of both larvae and adult 
ground-living beetles, Tenebrionidae, Staphylinidae, and a weevil, Blosyrus, also 
fragments of ants. 

Systematic Notes 

The (J from Sa da Bandeira and the unsexed specimen from Leba are notably 
larger than the other two. Wing 104-106 ; bill 23, 23 ; tail 105, 11 1 ; against 
wing 96, 96 ; bill 21, 22 ; tail 99, 93 for the specimens from Leba sexed as male and 
female. Furthermore, there is some difference in tail pattern between the two 
larger birds and the two smaller ; in the larger the black on the outer edge of the 
outermost rectrices extends about 56 mm. from the tip ; in the smaller pair it 
extends 37-40 mm. from the tip. It seems probable that an error was made in sexing 
the " male " from Leba and that this difference in size and tail pattern between 
the two pairs may be due to sexual variation. 



132. Cossypha polioptera ? polioptera Reichenow 1892 : Lake Victoria. 

Specimens 

1 cJ 2 $ Luacano, 2nd Aug. 



43© THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 

Systematic Notes 

C. p. grimwoodi White, was described from the Mwinilunga district as having a 
clearer grey head top, and the back and rump less tawny, more olive brown than 
polioptera. With seasonal variation and a certain amount of foxing in old skins 
it is difficult to place these. They have the head and mantle darker than grimwoodi, 
and the mantle, except in one female which was salted, more olive than available 
specimens of polioptera which may possibly be slightly foxed. 

I therefore refer these tentatively to polioptera which extends across the continent 
to northern Angola : possibly the division between the two races may be found to 
lie along the Congo /Zambesi watershed. 

I 33- ( a ) Cossypha natalensis intensa Mearns 1932. 

(b) Cossypha natalensis larischi Meise 1958, p. 73 : Canzele, Cuanza Norte. 

Specimens 

(a) 1 (J 1 ? Luau R., 29th July : 1 imm. ? Luacano, 2nd Aug. (b) 1 <J 2 $ 1 imm. $ 
12 m. S. of Gabela, i7-i8th Sept. 

Systematic Notes 

The distinctiveness of the north Angola population and variation within C. n. 
intensa have been discussed (Hall 1958 : 154). 

138. (a) Erythropygia leucophrys munda (Cabanis) 1880 : Malanje. 

(b) Erythropygia leucophrys ovamboensis Neumann 1920 : Ombongo,Ovam- 
poland. 

Specimens 

(a) 1 ? Quipungo, 29th Aug. (b) 1 $ E. of Cahinde, 6th Sept.: 1 ? 60 m. N. of 
Lobito on Novo Redondo road, 13th Sept. 

Systematic Notes 

It would seem that munda and ovamboensis are ecological races, munda being 
found in the woodlands and ovamboensis in the thorn country. Both the Quipungo 
and Cahinde birds were taken in areas where the two types of country meet, the 
first in a bush between a pocket of woodland and a pocket of thorn scrub, the second 
near the foot of the escarpment where stunted Brachystegia and mopane gives way 
to the semi-desert. This latter specimen has also some intermediate characters being 
more heavily streaked than is usual for ovamboensis, but is considerably paler above 
and less washed with rufous below than munda. 

139. Erythropygia leucosticta reichenowi Hartert 1907 : Canhoca, Cuanza Sul. 

Specimen 

1 imm. § Chingoroi, 9th Sept. 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 431 

Field Notes 

This elusive scrub robin was never seen, the single specimen being collected in a 
net set in slender, very close-growing trees with very little undergrowth, near the 
stream in the forest. 

Systematic Notes 

This is the most southern record for the species and I believe the first known 
female of this form. It was compared with two males from northern Angola borrowed 
from Chicago and was found to be a good match, only a trifle paler on the mantle. 
It still has some young feathers with rufous edges in the secondaries and wing-coverts. 

141. Acrocephalus baeticatus cinnamomeus Reichenow 1908. 

Specimen 

1 ? Chingoroi, 8th Sept. 

Systematic Notes 

The three races of Reed Warbler, baeticatus (Vieillot), cinnamomeus and suahelicus 
Grote, are, at best, only distinguishable in series. This specimen, in moult, is un- 
satisfactory for comparison but has been identified as cinnamomeus by Mr. C. M. N. 
White, who is preparing a revision of the species. 

143. Bradypterus tnariae boultoni Chapin, Ann. Cam. Mus. 31, p. 1, 1948 : 
Mombolo, Cuanza Sul. 

Specimens 

1 (J 2 ? Mt. Moco 7,000 feet, i6-23rd Aug. 

Field Notes 

This Forest Warbler was quite plentiful in the pockets of forest, as its voice, a 
loud " chep, chep " frequently repeated, was heard throughout the day, even when 
other birds were silent. It was not often seen, and always in thick undergrowth and 
very near the ground. The stomach contained crickets and other insects. 

142. Seicercus laurae laurae Boulton 1931 : Mt. Moco. 

Specimens 

2(j2?i Mt. Moco 7,000 feet, I9~22nd Aug. (1 in spirit.) 

Field Notes 

Two of these were a pair found in thickish forest, the female carrying nesting 
material of feathers. Another was one of a party of three or four feeding in foliage 
about ten feet from the ground. The food was found to be larvae and insect fragments. 

zool. 6, 7. 26 



432 THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 

145. (a) Calamocichla gracilirostris winterbottomi White 1947 : Manyinga R., 
Macondo dist., Moxico. 

(b) Calamocichla gracilirostris cunenensis Hartert 1903 : Cunene R., 
S. Angola. 

Specimens 

(a) 2 ■ (J 2 ? Lake Dilolo, 3~5th Aug. Wings <J 72, 74, ? 68, 69. (b) 1 ? Leba, 30th 
Aug. Wing 69 mm. 

Systematic Notes 

I have followed Chapin (1953 : 441) in assigning both these forms of Swamp 
Warbler to C. gracilirostris (Hartlaub). The birds from Lake Dilolo answer the des- 
cription of winterbottomi but are a little larger, the type series having wings of 67-71 
mm. The Leba specimen is not strikingly different but is a little paler on the back 
and is more washed with buff on the flanks and thighs. 

147. Calamonastes simplex huilae Meise 1958, p. 72 : Huila. 

Specimens 

1 $ Mt. Moco 6,000 feet, 24th Aug.: 1 $ Vila Flor, 26th Aug. Wings 63, 64 : 
bill 14 ; tail 46, 48 mm. 

Systematic Notes 

These Wren-warblers answer the description of Meise's new race, and the characters 
of these and some specimens from Vila da Ponte, collected by the Visser-Transvaal 
Museum, had already been observed by Dr. Rudebeck and myself before the publica- 
tion of Meise's paper. 

149. Apalis rufogularis brauni Stresemann 1934 : Roca Congulu, Cuanza Sul. 

Specimens 

1 cJ 2 juv. ? 12 m. S. of Gabela, i6-i8th Sept. 

Field Notes 

The adult was collected in the very top of a tall tree by the river ; the others 
were in the escarpment forest, one being a member of a family party of four in 
thickish trees about ten feet from the ground. 

152. Sylvietta r. ruficapilla Bocage 1877 : Caconda, Huila. 

Specimens 

1 $ Lake Dilolo, 5th Aug.: 1 (J 15 m. N. of Vila Flor, 25th Aug.: 1 $ Caconda, 
28th Aug. 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 433 

Systematic Notes 

All three specimens vary in the extent of rufous on the neck. The female from 
Caconda has none and has the thighs also grey ; the male from Vila Flor and a pair 
from Caconda in the British Museum have a narrow half circle of pale rufous on the 
neck and pale rufous thighs. The female from Lake Dilolo has a dark rufous patch 
on the neck and dark rufous thighs. It also has the white of the lores very much 
reduced, but in all other respects matches in colouring with the topotypical birds. 
On the limited material available it is not possible to judge the significance of this 
variation. 

154. (a) Sylvietta rufescens ansorgei Hartert 1907 : Huxe, Benguela. 

(b) Sylvietta rufescens mossamedes Meise 1958, p. 71 : 25 m. S. of Jau, 
Mocamedes. 

Specimens 

(a) 1 (J 1 $ 12-15 m. SE. of Benguela, n-i2th Sept. (b) 1 Leba, 31st Aug. 

Systematic Notes 

The possiblity of Sylvietta rufescens and 5. ansorgei being conspecific was suggested 
by Macdonald & Hall (1957 : 28) when a specimen from Ohopoho, Kaokoveld, 
was discussed. This specimen showed some intermediate characters, as does the 
unsexed specimen from Leba. Further similar specimens were examined by Meise 
who has given them the name mossamedes. The Ohopoho bird is in the Transvaal 
Museum and has not been compared with the Leba bird, but it seems certain that 
it will also be found to be this race. 

155. (a) Eremomela icteropygialis puellula Grote 1929 : Catumbela, Benguela. 
(b) Eremomela icteropygialis polioxantha Sharpe 1883 : Swaziland. 

Specimens 

(a) 1 $ Cahinde, 5th Sept.: 1 £ 12 m. SE. of Benguela, 12th Sept. (b) 1 100 m. 
W. of Vila Luso, 8th Aug.: 1 Vouga, 10 or nth Aug. Wing (a) <$ 50, (b) £ 53 mm. 

Systematic Notes 

The series of puellula in the British Museum shows that birds from south Angola 
are rather darker as well as smaller than South West African specimens (wings 
of 7 c? 49-55 ; 6 ? 48-53 against 4 $ 54-57 ; 9 ? 52-57 mm.). 

The two specimens of polioxantha were unfortunately badly shot and preserved 
in spirit ; they have since been skinned but are not very satisfactory for comparison. 
This is unfortunate as they are important specimens showing that grey-backed 
birds are found well within the range of the green-backed E. salvadorii (see species 
158) supposed by some authors to be conspecific. There is, however, no question 



434 THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 

that any green on the mantle might have been lost in the spirit since these were 
grey when collected, the only green-backed Eremomela being obtained later at 
Vila Flor and recognized then as something new to our collection. 

The races puellula and polioxantha provide examples of strongly differentiated 
races specialized in thorn and Brachystegia country. 

157. Eremomela atricollis Bocage 1894. 

Specimens 

2 cJ 5 m. W. of Vila General Machado, 10th Aug.: 1 cJ 1 $ 15 m. N. of Vila Flor, 
25~26th Aug.: 1 ? 5 m. S. of Caconda, 28th Aug. 

Field Notes 

The Black-necked Eremomela was found in fairly tall and in stunted Brachystegia. 
It was noted to be very active, fossicking for insects with tit-like agility, often 
upside down. It was once found with a small party of E. scotops. 

158. Eremomela salvadorii Reichenow 1891 : Leopoldsville, Belgian Congo. 

Specimen 

1 $ 5 m. N. of Vila Flor, 26th Aug. 

Systematic Notes 

Specimens of the green-backed species in the British Museum have been obtained 
at Nova Lisboa, Vila Luso, both by Admiral Lynes, and now at Vila Flor. They 
establish it as sympatric with E. i. polioxantha and as a bird of bushes on the edge 
of woodlands, thus requiring very similar but possibly more open habitat. There 
seems no significant difference in colour between the Angola birds and some from 
Luluabourg in the Kasai, and I follow the authors in considering E. griseoflava lundae 
Grant & Praed, a synonym of E. salvadorii. 

159. (a) Camaroptera brevicaudata sharpei Zedlitz 191 1 : Damaraland. 

(b) Camaroptera brevicaudata subsp. (?) 

(c) Camaroptera brevicaudata harterti Zedlitz 1911 : Canhoca, Cuanza Sul. 

Specimens 

(a) 1 <$ Leba, 3rd Sept.: 1 <$ 1 juv. Chingoroi, 8th Sept.: (b) 1 <$ Luau R., 28th 
July (and one in spirit), (c) 1 $ Gabela 17th Sept. 

Systematic Notes 

The green-tailed form harterti with very white underparts is strikingly different 
from other races of C. brevicaudata, all of which have brown or grey-brown tails 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 435 

and are heavily washed with grey or buff on the chest. Its range is apparently 
restricted to localities on and below the escarpment from Ndala Tando and Luanda 
to Gabela and the specimens were probably all collected, as ours was at Gabela, 
in the escarpment or riverine forests. It is the more surprising therefore that other 
forms have been collected in similar forests at Chingoroi and Canzele, ours from 
Chingoroi being netted alongside such a typical forest bird as the Wattle-eye, 
Dyaphorophia concreta. The Canzele specimens in the Chicago Museum are those 
referred by Heinrich (1958 : 349) to C. b. harterti and which proved on examination 
to be C. b. tincta. 

It seems possible that harterti is the true ecological representative of C. brevicaudata 
in the escarpment forests, but that, on the northern and southern edges brown- 
tailed forms have intruded on its habitat, and it would not be surprising if harterti 
was eventually found as well at Canzele or Chingoroi. 

Among the brown-tailed races it has been found, as with many Cisticolas, that 
there is little or no seasonal variation in some of the tropical forms, but well-marked 
variation in the sub-tropical forms. C. b. sharpei of southern Africa (of which 
C. b. beirensis Roberts, is considered a synonym) is an example of the latter, non- 
breeding birds having mouse-brown backs and the underparts washed with buff, 
breeding birds having grey backs and greyish white underparts. North of sharpei 
on the east of the country is erlangeri Reichenow, of eastern Tanganyika, with little 
seasonal change and characterized by very white underparts and a greyish-brown 
back. North again is griseigula with the underparts strongly washed with grey. 
On the west, north of harterti, is the dark grey tincta (Cassin), with no seasonal dress. 
This is found in Gaboon, Cameroons and northern Angola, extending into the northern 
Congo where it intergrades with East African races (see Chapin 1955 : 317). 

However, in central Africa north of the recognizable limits of sharpei, west and 
south-west of erlangeri and griseigula, and south of tincta, there are specimens 
which represent an intermediate or several intermediate populations. These are 
similar above to erlangeri and griseigula in having grey-brown mantles, and below 
are closest to the breeding dress of sharpei but with slightly less grey on the throat 
and slightly washed with buff on the flanks. There seems to be little or no seasonal 
change. 

Specimens examined which show these characters are as follows : 

Angola : 1 ? Malanje, 24th Feb.; 1 $ Missao de Luz, Jan, coming into breeding 
condition ; 1 juv. <J Saurimo, Lunda dist. 20th Dec. (the two latter previously 
identified as harterti, Ibis, 1934 : p. 46). 1 (J Luau R., 28th July, not in breeding 
condition. 

Belgian Congo : 1 <$ Kayoyo (io° 30' S. 24 30' E.), 1st Sept. coming into breeding 
condition. 1 Kambove, 1st July. 

NW. Rhodesia : 1 $ Kabompo R., 9th Feb. 1 imm. $ Mwinilunga, 9th Jan. 

Tanganyika : 1 <J Lukolansala R. Western Prov. 20th Nov.; 1 (J Mt. Hanang, 
nth Feb.; 1 ? Usangu dist. 5th Nov.; 2 <J 2 ? 1 juv. <J Iringa uplands, 19th Jan. 
and 5th March, the March specimens post-breeding. 

It will be seen that this is a very inadequate series from widely scattered localities 
and without proper seasonal representation in any one place, and it may well be 



436 THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 

that adequate series will show that birds from these localities are not so homogenous 
as now appears. Until more material is available I do not feel that there is justifica- 
tion for giving them a name. 

The adults collected at Chingoroi and Leba are typical of sharpei in non-breeding 
dress. 

162. Cisticola chiniana near j rater. 

Specimen 

1 (<£ on size) Leba, 31st Aug. 

Systematic Notes 

This specimen is rather darker than f rater Reichenow, from Damaraland, as are 
three collected by Lynes in 1931 from the Que R. (14 30' S., 14 30' E.) which he 
labelled as " near f rater." 

181. Hirundo abyssinica unitatis Sclater & Praed 1918. 

Specimen 

1 Mt. Moco 7,000 feet, 17th Aug. 

Field Notes 

This Striped Swallow was observed to be apparently nesting in the cliff face over 
a waterfall in the mountain forest. This was unexpected as Chapin (1953 : 761) 
notes that in the Congo it is wanting on all the higher mountains, especially those 
that are forested. 

190. Ptynoprogne fuligula near rufigula. 

Specimen 

1 Mt. Moco, 22nd Aug. Wing 121 mm. 

Systematic Notes 

Chapin (1953 : 744) suggests that Rock Martins from the central highlands of 
Angola are intermediate between rufigula (Fisher & Reichenow) of East Africa and 
the pale anderssoni (Sharpe & Wyatt), of South West Africa. This specimen is 
true rufigula in colour, but is larger, as is anderssoni. 

192. Petrochelidon rufigula (Bocage) 1878. 

Specimens 

3 (J 6 $ 20 m. W. of Vila General Machado, 10th Aug. (and one in spirit) ; 1 $ 2 £ 
1 Nova Lisboa, I2-I3th Aug. 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 437 

Field Notes 

This cliff swallow was found breeding in a colony under a concrete road bridge. 
There were about two hundred and fifty nests massed together ; each was a large 
half sphere approximately 9 inches wide and 4 inches deep with an entrance spout 
varying greatly in construction but always very wide, about 2 inches across. The 
nest was lined with fine grass, no feathers. A clutch of two eggs was taken from one 
nest and a single egg from another. Several pairs of Micropus caffer were parasitizing 
some of the nests. In the Lower Congo P. rufigula was found breeding in April 
(Chapin 1953 : 773). 

Systematic Notes 

Chapin notes that P. rufigula is closely allied to P. preussi (Reichenow). Their 
affinities are further stressed by the eggs, which were examined by Dr. W. Serle, 
to whom I am indebted for the following notes. 

Eggs : (Notes made by Dr. W. Serle, 31.xii.58). 

" Eggs ovate, slightly glossy, smooth surface. Ground colour white tinged creamy 
pink. Spotted and blotched with reddish-brown and chestnut, brown primary and 
ashy violet secondary markings, both primary and secondary markings being densest 
about the large end and where they coalesce to form a ring or cup. One egg shows 
a few black hair lines. They measure 18-2 X 12-9 ; 19-5 X 13 ; 19-3 X 13-1 mm. 

"In appearance they closely resemble one type of Petrochelidon preussi (Reichenow), 
from Nigeria, and on the criterion of the egg (size, shape, ground colour, markings, 
texture etc.) there is no reason why Petrochelidon rufigula and P. preussi should 
not be conspecific. On the other hand, these three P. rufigula eggs do not resemble 
Petrochelidon spilodera eggs from South Africa (of which I have seen many), nor do 
they resemble the eggs of Petrochelidon fuliginosa (Chapin) of the Cameroons. 

"The eggs of P. preussi and P. rufigula are distinctive. I do not know any other 
African swallows' eggs that resemble them/' 

199. Prionops gabela Rand, Fieldiana 39, p. 43, 1957 : 15 km. S. of Gabela. 

Specimens 

1 imm. $ 12 m. S. of Gabela, 20th Sept.: 1 $ 40 m. S. of Mumbondo on Gabela- 
Muxima road, 20th Sept. 

Field Notes 

The first specimen of this new Helmet Shrike was obtained by Heinrich in " a 
little region of tangled tropical second growth along the ridge of a higher mountain 
range." From his subsequent paper (1958 : 128) it seems that he was camped a 
few miles further up the Cuvo River, nearer Assango, than we were. Our first 
specimen was obtained also on a ridge above the coffee forest where the trees were 
less thick. The second was found in the country below the escarpment ill the trees 



438 THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 

of a dry thicket by the road in an area of mixed thickets and cultivation. Both 
were members of small parties. 

Systematic Notes 

These are the first known females of this species. They answer the description 
of the male except that both have some white on the 2nd-6th primaries. This 
white is confined to very small patches on the inner web ; it is slightly more extensive 
in the adult, and in the young bird is asymmetrical. It is in no case comparable to 
the large patches in P. retzii. 

Signs of immaturity in the Gabela specimen are found in a few brown feathers, 
very worn, among the fresh black feathers of the head, and in the colours of the 
iris, bill and legs as compared with the adult. 

Imm. <j>, Gabela Ad. $, Mumbondo 

Iris . . . Yellow . Yellow with red outer 

ring. 
Bill . . . Red base, orange tip . Red, orange tip to lower 

mandible. 
(In the dried skin the orange of the greater part of the lower mandible of the 

young bird is very noticeable.) 
Legs . . . Bright orange-red . Cherry-red. 

Eye wattle . . Red . Red. 

204. Laniarius aethiopicus major (Hartlaub) 1865 : Elminia, Gold Coast. 

Specimens 

1 imm. ^ Luau R., 31st July : 1 <J Lake Dilolo, 5th Aug. 

Field and Systematic Notes 
See next species. 

206. Laniarius bicolor guttatus (Hartlaub) 1865 : Benguela. 

Specimens 

1 cJ 2 $ 1 imm. $ Leba, 31st Aug.-i Sept.: 1 juv. Chingoroi, 8th Sept. 

Field Notes 

The relationships and habitats of the Boubou Shrikes were discussed (Hall, 1954), 
and it was suggested that the different species were more restricted in their choice 
of habitat in the areas where two species were near neighbours. It was particularly 
interesting therefore to have my first experience of them in the field in Angola 
where L. aethiopicus and L. bicolor adjoin. Disappointingly little was seen of these 
shrikes in the critical central areas, the only specimen shot being one in the foothills 
of Mt. Moco which was never recovered from a thick reed-bed in the valley of a 
stream. 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 439 

However, in the eastern districts L. aethiopicus was found to be an inhabitant 
of scrubby bushes in the Brachystegia woodlands, as it is in south-western Northern 
Rhodesia and the Caprivi Strip, where it occurs alongside L. b. sticturus Finsch & 
Hartlaub. 

In the west L. bicolor was collected at Leba in thick cover by the stream or on 
the hillside just above, at Chingoroi in cover by the stream, and from a dry tangled 
thicket in the grass and thorn country above the sea near Novo Redondo. These 
different habitats in the extreme west conform to the suggestion that the habitats 
are less specialized away from competition with the related species. 

Systematic Notes 

The adults of the two species are readily distinguished by the relative whiteness 
of the underparts, and the observations made in the previous paper on the eye- 
colour of the two species, was borne out by our specimens ; the adult of L. aethiopicus 
being noted as having the iris " rich dark brown " and those of L. bicolor having it 
" dark brown." 

The young birds are harder to distinguish since the pinky tinge on the breast of 
aethiopicus and the pure white of bicolor are replaced in the young by a buffy tinge 
in both species. But, as with the adults, when the feathers of the breast are lifted 
it will be seen that in aethiopicus they become more tinted with colour towards 
the base, which is black, whereas in bicolor they become whiter towards the black 
base. 

Leba lies on the borders of two races, L. b. guttatus (smaller with two secondaries 
usually edged white) and L. b. sticturus (larger with three secondaries usually edged 
white). Two Leba birds have a narrow edge on the third secondary but in respect 
of size (wings 92-96) are clearly best referred to guttatus. 

211. Tchagra s. senegala (Linnaeus) 1766 : Senegal. 

Specimens 

i(Ji? Luau R., 27th July : 1 $ 40 m. W. of Texeira de Sousa, 31st July : 1 $ 
Mt. Moco 6,000 feet, 18th Aug. 

Systematic Notes 

Two of these Tchagras are just finishing moult, the other two just starting, so 
they illustrate well the extremes of seasonal variation. The two fresh birds were 
found to match well with some fresh specimens from West Africa. I therefore do 
not recognize rufofusca (Neumann), described from north Angola as distinct from 
the nominate race. 

214. Chlorophoneus multicolor batesi Sharpe 1908 : R. Ja, Cameroons. 

Specimens 

1 ^ 1 imm. ^ Gabela, 16th Sept. 



440 THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 

Systematic Notes 

Moreau & Southern (1958) discuss colour variation in this and the following 
species of bush-shrike. The adult bird is in their " Yellow D " phase (p. 316) as are 
all others that were examined from north Angola : the young bird has yellow, not 
orange, underparts, light tips to the wing-coverts, light edges to the gape, and was 
noted as having a dark brown, not a deep purple, iris. 

In their paper no mention was made of differences in size between Angola and 
Cameroon birds but later Moreau, examining specimens with me, found that those 
from Angola, including some borrowed from Chicago, have smaller bills. The bills 
of two from Gabela are particularly small (19, 20 mm.), four from further north 
in Angola have them slightly larger (20-21 mm.), while the bill in Cameroon birds 
is 21-23 mm. and deeper in proportion. 

There is also a tendency for Angola birds to have the grey of the crown and hind 
neck more sharply demarcated from the mantle than in Cameroon birds, but this is 
not constant. 

215. Chlorophoneus nigrifrons manningi (Shelley) 1899. 

Specimen 

1 imm. <J Luau R., 29th July. 

Field Notes 

Moreau and Southern (1958 : 308) found that the population of the Blackfronted 
Bush-Shrike in Northern Rhodesia was less restricted ecologically than others. 
This was confirmed by our specimen which was found not far from the river in light 
Brachystegia woodland, and not in forest, as is more usual. 

217. Telephorus viridis (Vieillot) 1817. 

Specimens 

i(Ji? Chingoroi, 9-ioth Sept. Wing <$ 90, ? 86 mm. 

Field Notes 

Both of Perrin's Bush-Shrikes were collected, one being netted, in thick undergrowth 
of the forest. The call was described as " not unlike a francolin call, cut off very 
short. When hopping about utters a sharp but muted ' chunk . . . chunk . . . 
chunk.' " 

Systematic Notes 

The race T. v. vieiriae White, was described from eastern Angola on slight differences 
in size and colour in comparison with the typical race, in particular on evidence 
of sexual dimorphism not believed to exist in birds of western Angola. Examination 
of the series in the British Museum shows that there are differences between the 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 441 

sexes in both populations, the female always having the red and green of the throat 
and underparts paler and duller. Furthermore, these two birds are larger than those 
previously recorded from the west and similar in size to White's series. I can therefore 
see no grounds for recognizing T. v. vieiriae. 

218. Malaconotus hypopyrrhus interpositus Hartert 191 1 : 40 m. W. of Baraka, 
Lake Tanganyika. 

Specimens 

1 <J Luau R., 29th July : 1 $ 5 m. W. of Munhango, 9th Aug. 

Systematic Notes 

The taxonomy of the Grey-headed Bush-Shrike has been confused by differences 
of opinion on the correct use of blanchoti Stephens, poliocephalus (Lichtenstein), 
or hypopyrrhus (Hartlaub), as the specific name, and on the importance of an orange 
wash on the underparts, and of a white circle round the eye as taxonomic characters. 

I follow Grant & Mackworth-Praed (1958 : 19) in using hypopyrrhus as the specific 
name, and follow Chapin (1954 : 42) in recognizing geographical as well as individual 
variation in the amount of orange wash on the underparts. I do not however, agree 
with Chapin's treatment of the Angola birds, all of which he calls monteiri, distin- 
guishing them from interpositus of Rhodesia by the lack of any orange wash below 
on the majority of specimens. The white circle round the eye, characteristic of the 
type of monteiri (Sharpe) , from near Luanda, and the type of perspicillatus (Reichenow) 
from Cameroon Mt. he considers an abnormal character of no taxonomic significance. 

Other specimens with this character are from Dondo and Ndala Tando (in the 
British Museum), Gabela and " Bucaso " (in Chicago). With the exception of 
Cameroon Mt. and " Bucaso ", which has not been located, all these places are along 
or below the escarpment in north-western Angola, and all have riverine and escarp- 
ment forest rather than woodland. All other specimens examined from Angola in 
Chicago or London have been taken from above the escarpment in more predomi- 
nantly woodland areas, and have the white on the face variable and restricted 
to areas between the eye and the bill, or on the lores. 

There may thus be an ecological as well as morphological difference between the 
birds with the white ring and those without and, in my opinion, the name monteiri 
should be used only for those with this character. The race therefore has a restricted 
range in north-west Angola and possibly a discontinuous distribution north to 
Cameroon Mt. (see Serle 1954 : 72). The colour of the underparts of monteiri is 
variable, the Dondo and Ndala Tando specimens have an orange wash, the others 
being clear yellow. 

All other specimens from Angola, without the eye-ring, I would include with 
interpositus, at least for the present, though noting that the orange wash commonly 
found in specimens in the east of the range becomes less frequent and less pronounced 
in specimens in the west. 

In studying these shrikes I am indebted to Dr. A. L. Rand for a copy of his notes 
on the species and details of specimens in Chicago. 



442 THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 

225. Neocichla g. gutturalis Bocage 1871 : Huila. 

Specimens 

2 (J 2 $ 48 m. SW. of Caconda, 28th Aug. (one in spirit). 

Field Notes 

For some distance along the Caconda-Quipungo road these Babbler-Starlings 
were common, flying up from the roadside into bushes and low branches when 
disturbed by the car. This was in the area of leafless Brae hystegia- type woodland 
mentioned in the discussion on vegetation. 

Systematic Notes 

These are the first specimens of the nominate race in the British Museum. In 
comparison with N. g. angusta Friedmann, they seem to have the head and throat 
a paler, clearer grey as well as having larger white tips to the tail. 

228. (a) Lamprotornis m. mevesii Wahlberg 1857 : Okavango, N. Bechuanaland. 

(b) Lamprotornis mevesii benguelensis Shelley 1906 : Capangombe, S. 
Angola. (= L. m. purpureas auct.) 

Specimens 

(a) 1 (J 25 m. E. of Sa da Bandeira 4,900 feet, 27th Aug.: 1 (J 10 m. E. of Cahinde 
3,000 feet, 6th Sept. (b) 1 $ 15 m. S. of Quilengues, 3,900 feet, 7th Sept. 

Systematic and Field Notes 

The tw T o races of Long-tailed Starling are clearly differentiated in the blue or 
bronze sheen of the whole plumage and none of the specimens examined show signs 
of intergradation. The boundary between them seems interesting following the 
discussion on the three avifaunal zones in Angola, for the bronzy L. m. benguelensis 
seems to be restricted to what might be termed the southernmost tip of the Escarp- 
ment Zone, a narrow tongue of rather richer vegetation lying between one thousand 
and three thousand five hundred feet with the coastal semi-desert on the west and 
the Brachystegia woodland on the east. Specimens of benguelensis have been obtained 
at Catengue, near Chingoroi, Quilengues, Vila Arriaga and Capangombe. Quilengues 
lies more than halfway along the Sa da Bandeira to Chingoroi road which descends 
gradually with the vegetation becoming progressively richer and more varied 
northwards. Vila Arriago and Capangombe lie in fertile pockets at the base of the 
Chela escarpment but bordered westwards by the desert. 

L. m. mevesii on the other hand has a widespread distribution in the Mopane 
belt from Nyasaland westwards, and it is confined in Angola to the less arid parts 
of the Acacia Zone. It has been collected at Jau, Humbe, Chipopia, Mulondo and 
Mupa. The specimen collected east of Cahinde was one of a number seen on the road 
down the escarpment, where there was a light covering of mixed bush and stunted 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 443 

trees including Mopane. Although Cahinde lies like Capangombe at the foot of 
the escarpment, and is only forty miles south, it has not the same rich vegetation 
and the associations of all species collected there are with the Acacia Zone. 

It should be noted that some of the specimens collected by Monard (1954 : 83, 84) 
were examined with Captain Grant in 1957 through the kindness of the Director of 
the Musee d'Histoire Naturelle, Chaux de Fonds. Those from Mupa and Mulondo 
identified as purpureas were found to be L. m. mevesii, and samples of those identified 
as mevesii were found to be L. australis. 

231. (a) Zosterops senegalensis quanzae de Schauensee 1932 : Upper Cuanza R. 

(b) Zosterops senegalensis heinrichi Meise 1958 : Canzele, Cuanza Norte. 

(c) Zosterops senegalensis quanzae % anderssoni Shelley. 

Specimens 

(a) 1 Mt. Moco 6,500 feet, 15th Aug.: 1 $ 25 m. N. of Quipeia, 14 Aug. (b) 1 $ 
12 m. S. of Gabela, 19th Sept.: 1 <J 28 m. N. of Vila Luso, 8th Aug. (c) 1 <J Leba, 
30th Aug. 

Systematic Notes 

Identifications have been made on the basis of Moreau's paper (1957). These 
specimens have interest in filling some of the gaps that were then noted. The Moco 
and Quipeio birds have the characteristic dull colouring and large size (wing 65, 62) 
of typical quanzae. The Gabela and Vila Luso birds are a shade brighter, particularly 
on the throat and tail-coverts, and the Gabela bird on the forehead. They are close to, 
but a little more olive than, the Ndala Tando birds quoted by Moreau which have 
since been named heinrichi. 

The Leba bird is altogether brighter below, matching in this respect a topotypical 
anderssoni from Elephant Vley, but being rather darker above and larger (wing 61). 

232. Nectarinia kilimensis gadowi Bocage 1893 : Galanga, Benguela dist. 

Specimens 

1 (J Vila Flor, 13th Aug.: 9 $ 1 imm. <J 1 <j> Mt. Moco 6,000-7,000 feet, i5~2ist 
Aug.: 1 (J 2 imm. c? 1 ? 1 imm. Nova Lisboa, 13-14^ Aug. 

Field Notes 

This sunbird was common on Mt. Moco at all levels, feeding on Erythrina and 
Loranthus. Stomach contents included spiders, Coleoptera, Diptera, and hover-flies. 
Its call was noted as being exactly like that of the nominate race. 

233. Nectarinia bocagei Shelley 1879 : Caconda, Huila dist. 

Specimens 

1 $ 3 imm. $ near Vila Flor, I3~i4th Aug.: 1 $ near Luimbale, 25th Aug. 



444 THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 

Field Notes 

Bocage's Sunbird was rare. It was once found feeding on Erythrina with Cinnyris 
oustaleti and Chalcomitra senegalensis, and once on red Loranthus in Brachystegia 
woodland. 

237. Cinnyris o. oustaleti (Bocage) 1878 : Caconda, Huila dist. 

Specimens 

1 $ 1 imm. $ Nova Lisboa, 12th Aug.: 1 J 43 m. W. of Vouga, 12th Aug.: 2 <£ 
1 imm. ? near Vila Flor, I3~26th Aug.: 4 (J 1 juv. Mt. Moco 6,000-6,500 feet, 
i5~24th Aug. 1 imm. $ near Luimbale, 25th Aug. 

Field Notes 

Found feeding on orange-red gladiolis in the Brachystegia woodland on Moco, 
and on Albizia trees at Luimbale. Its food contained seeds, spiders, beetles and 
Lepidoptera larvae. 

Systematic Notes 

The seven males, including one in which the skull was not fully ossified, are fairly 
uniform in appearance with a few metallic feathers on the head and mantle and the 
metallic plumage below confined to the centre of the throat and chest. In four of 
the specimens sheath feathers of metallic plumage were found on the chest and mantle ; 
in three, including the immature bird, the wings and tails are in moult, in the others 
they are fresh. The adult female is moulting wings and mantle. Williams (1955) 
discusses the moults and plumages of this species, finding that the moult from 
eclipse to metallic plumage is complete, including wings and tail, but that the wings 
and tail are not moulted after breeding. It would seem therefore that the males 
collected are moulting into breeding dress. 

The finding of these males concurrently with young birds with skulls 10 % or 

20 % ossified suggests a protracted breeding season. 

The identification of the juvenile from Moco is not certain, no comparable speci- 
mens being available. 



246. (a) Anthreptes collaris somereni Chapin 1949 : Anda, Lake Azingo, 
Gaboon. 

(b) A.c. phillipsi White 1950 : Lofu R., N. Rhodesia. 

Specimens 

(a) 1 cJ 1 $ 12 m. S. of Gabela, I5~i6th Sept. (b) 1 <$ 1 $ Luau R., 27-29A July ; 
1 imm. (J 1 imm. $ Lake Dilolo, 5th Aug. 1 <J 45 m. W. of Vila Luso, 8th Aug. 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 445 

Systematic Notes 

The adult female from the Luau River has the grey throat and chest characteristic 
of phillipsi well marked. The Gabela pair have been identified as somereni on 
locality. In colour they seem indistinguishable from zambesiana (Shelley), though 
they are smaller. Both are bright clear yellow below, without much olive wash on 
the flanks, and the female has a pale whitish-yellow throat. Comparison between 
somereni and zambesiana is a little difficult on available skins in the British Museum 
since most of those of somereni are small, rather pinched skins which exaggerate the 
dusky flanks and hide the clear yellow of the tummy ; most of those of zambesiana, 
and the Gabela pair, are well filled-out skins which show off the yellow. It seems 
that there may not be such clear-cut differences between the two races as appears 
from this series. 

252. Passer griseus griseus (Vieillot) 1817. 

Specimen 

1 $ 12 m. S. of Gabela, 17th Sept. Wing 78, bill 15 mm. 

Field and Systematic Notes 
See next Species. 

253. Passer diffusus georgicus Reichenow 1904 : Damaraland. 

Specimen 

1 $ Leba, 31st Aug. Wing 78, bill 13*5 mm. 

Field and Systematic Notes 

The relationship between the two forms of grey-headed sparrow found in coastal 
Angola has been discussed by Benson (1956) and by White & Moreau (1958), 
and the authors differ in their views as to whether the two forms can be regarded 
as conspecific. The two specimens collected conform with Benson's conclusions 
on the ecological segregation of the forms in western Angola, since the male griseus 
was obtained in the vicinity of native huts in a clearing in the coffee forest, while 
the female diffusus was found in a mountain valley some distance from habitation. 

The Leba bird, like others of diffusus from Angola, noted by White & Moreau, is 
small and its bill is horn-coloured, paler below, with a yellow gape. The Gabela 
griseus is also short-winged in comparison with other Angola specimens noted, but 
has a heavy bill which is black. 

257. Ploceus temporalis (Bocage) 1880. 

Specimens 

1 cJ 1 (with nest), Saiangikilo R., 40 m. W. of Munhango, 9th Aug. Wing 81-84 ; 
bill 19 ; tail 44-45 mm. Bill black ; legs pale reddish-horn ; iris cream. 



446 THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 

Field Notes 

This rare weaver was only encountered once at a place where the road crossed the 
river in the centre of an open dambo. The nest, which is unused, was hung in 
elephant grass beside the river. (Plate 5 B, and 6). 

Nest. The nest has been examined by Professor and Mrs. N. E. Collias of the 
University of California, particularly in respect of the possible relationship of 
P. temporalis with P. capensis. I am grateful to them for the following comments, 
and for the photograph. 

" The nest is typically ploceine in its use of green, more or less flexible and inter- 
woven materials. It is a typical Textor (Ploceus) nest in its ovoidal shape, pensile 
attachment, ventral entrance, and presence of a special ceiling put in before the 
nest is lined below. The nest you sent us is unlined and apparently never contained 
eggs. It belongs to the same size class as nests of Textor cucullatus, T. spekei and 
T. capensis and not to the size class of nests of the smaller Textor s. 

" The nest has one specialized feature, unique so far as we know, among nests of 
the Plocinae. This is the extensive use, for construction of the outer shell, of grass 
culms from which the flowering tops have apparently been snipped off. 

" But on the whole the nest is a primitive one for a Textor, in various ways : loose, 
untidy weaving, use of grass stems in the outer shell, loose fastenings of individual 
pieces of nest material as well as loose attachments to the support, irregular ceiling 
of narrow sections of leaf blade, absence of longitudinally-stripped pieces of leaf 
blade, and in the occasional presence in the ceiling of grass heads (identified as 
Pennisetum for us by our Botany Department) . 

" In many of these features the nest resembles most closely that of Textor spekei 
of East Africa. However, the poor ceiling of narrow bits of grass blade seemed to 
us to be most like that of a young male T. cucullatus. In the presence of grass heads 
in the ceiling it resembles the nests of Othyphantes, Textor rubiginosa and Textor 
xanthops ; however, it lacks the abundance of grassheads that is found in the ceiling 
of these weavers. 

" The nest of Ploceus temporalis does not seem to us to closely resemble the one 
nest of Textor capensis that we have been able to examine closely . . . This 
capensis nest had the outer shell built of grass leaf strips and palm strips rather 
than grass stems. It had the ceiling made not only of broad (1 cm.) grass-leaf 
sections but also of pine needles and cypress branchlets. We recognize, of course, 
that a nest is only one clue to the taxonomic affinities of birds." 

Systematic Notes 

Having regard to Professor Collias's comments on the differences between the 
nests of P. capensis and P. temporalis as well as to the differences in the bills of the 
two forms, I do not feel that temporalis should be regarded as a race of capensis, 
at least until more is known of its field characters. 

The unsexed specimen is a good match with the plate in Shelley (1905 : pi. 41) ; 
both from its size and colour there is little doubt that it is also a male. The other 
specimen differs in having the olive of the ear-coverts extending on to the chin 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 447 

and centre of the throat making a tongue of dark colour sharply defined from the 
yellow of the underparts. 

258. Ploceus xanthops (Hartlaub) 1862: "Angola" = Lower Cuanza River 
(see below). 

Specimens 

1 (J 2 $ Mt. Moco 6,000-7,000 feet, I5~22nd Aug. 2 (J Leba, 30th Aug.~3rd Sept. 
1 ? Chingoroi, 8th Sept. 

Systematic Notes 

The type locality of Ploceus xanthops is given only as " Angola." The original 
description however is among notes on a small collection made by Monteiro of 
birds from the coastal areas of north Angola. Definite localities are given for other 
species, either Cuanza River, Cambambe or Massangano, while the weaver is noted 
as being " common everywhere . . . particularly over water." It seems that 
the type locality can therefore be usefully restricted to the Lower Cuanza River 
and specimens collected from Luanda, Dondo and Vila Salazar can be considered 
as typical. 

This is important since examination of skins in the British Museum, and some 
from Chicago, show that there is considerable geographical variation within Angola, 
particularly in the females. The variation is particularly interesting since it follows 
exactly that found in the Zosterops of Angola. Specimens of typical xanthops in the 
north are rather a yellow green matching the Zosterops described by Moreau (1957 : 
372) as intermediates between Z. s. kasaica and Z. s. quanzae and since named 
as heinrichi Meise (1958 : 80). Those of the Benguela highlands are a darker and 
duller green, matching Z. s. quanzae, while those of the south are brighter, more 
washed with yellow as is Z. s. anderssoni. 

When specimens of Ploceus xanthops from the rest of Africa were examined, 
including some borrowed from Pretoria, it was found that this variation in Angola 
represented the extremes of variation in colour found throughout the continent. 
The populations of north-eastern Africa, which have sometimes been separated as 
camburni (Sharpe), are brighter and more yellow than typical xanthops but similar 
to the southern birds. The populations of southern Africa, jamesoni (Sharpe), are 
very slightly greener and duller than typical xanthops with usually slightly less 
suggestion of dark streaks on the mantle but not so green or dull as the highland 
birds of Angola. Most specimens from central Africa match typical xanthops, but 
occasional birds, nearly all of which are from highlands of either southern Tangan- 
yika or Nyasaland, are dull and green, very close to the highland birds of Angola. 
This is again consistent with Moreau's findings on dark populations of Zosterops at 
higher altitudes (loc. cit.: 370). 

In view of the sporadic distribution of darker and brighter populations it is difficult 
to see how any races can be recognized on colour differences. In size the wing 
measurements of all males examined are between 86 and 96 mm. and all females 

zool. 6, 7. 27 



44« 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 



between 80 and 92 mm. with the exception of three from Maun and Lake Ngami, 
the male of which has a wing of 102 and the females 92 and 94. The name maunensis 
(Roberts), was given to birds from Maun and from the Chobe River ; the latter are 
no larger than typical xanthous so that if maunensis is to be considered distinct 
from xanthops it must be restricted to Lake Ngami. In my opinion it is hardly 
consistent in an admittedly variable species to single out the characters of this one 
population for recognition by name. 

The following table shows that there are other local variations in size but none of 
any significance. An attempt has also been made in the table to indicate the colour 
variation in the populations though it must be appreciated that there is also con- 
siderable individual variation particularly in the amount of yellow on the forehead 
of males. 

Table. — Populations of Ploceus xanthops 









Wing length 

A , 




Area 


f 

6* 




Colour 


(1) Angola, north 


12 $ 


7? • 


86-93 


80-85 


Yellow green. 


(2) „ highlands . 


36* 


5$ • 


90-91 


82-89 


Dull green. 


(3) „ south 


10 <J 


6? . 


88-94 


81-90 


Bright yellow green. 


(4) N. Rhodesia and S. 


10 q* 


7? • 


92-96 


84-90 


As (1) or between (1) 


Belgian Congo 










and (3). 


(5) Nyasaland and 


27 6* 


26 $ 


88-95 


84-90 


As (1), occasionally 


northern P.E.A. 










near (2). 


(6)S. Tanganyika 


26* 


5? 


90-92 


86-88 


. Slightly paler than 


(highlands) 










(2). 


(7) Rest of Tanganyika 


96* 


3? 


90-97 


88-89 


• As (3). 


(8) Kenya and Uganda 


21 6* 


12 $ 


91-97 


86-92 


. As (3), occasionally 
near (2). 


(9) P.E.A. and S. Rho- 


56* 


3$ 


89-94 


84-92 


. Between (1) and (2), 


desia 










slightly less streaky. 


(10) Southern Africa 


46* 


4? 


90-94 


84-91 


. As (9). 


(11) Chobe R. and Cap- 


36* 


4? 


93-94 


85-89 


. As (9). 


nvi 
(12) Lake Ngami . 


16* 


2? 


102 


92-94 


. As (1). 



264. Euplectes aurea Gmelin 1789. 

Specimen 

1 <J 12 m. E. of Benguela, 12th Sept. 

Systematic Notes 

This male is in very fresh non-breeding dress ; it appears identical in plumage with 
females. 



Cryptospiza r. reichenovii (Hartlaub) 1874 

Specimens 

1 cJ 1 £ 1 imm. (J 12 m. S. of Gabela, I7~i8th Sept. 



THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 449 

Field Notes 

Heinrich also found the Crimson-wing in the forest of Gabela but it has not been 
recorded from elsewhere in Angola. That it should be found here, and not in the 
montane forest on Moco, is surprising since it is more usually a mountain bird, 
being found on Cameroon Mountain and the highlands from Ruwenzori to Mashona- 
land, though descending to the coastal plain at Amani. 

Systematic Notes 

Chapin (1954 : 486) notes the difficulties of distinguishing between the races of 
this species. From examination of the series in the British Museum the only geogra- 
phical variation apparent is that the breasts and abdomens of all specimens from the 
Cameroons, Ruwenzori, Mufumbiro Volcanoes and Angola are more washed with 
orange, less green, than in specimens from Tanganyika, Nyasaland and Portuguese 
East Africa. I would recognize therefore only the two races C. r. reichenovii and 
C. r. australis Shelley. 



281. Clytospiza c. cinereovinacea (Sousa) 1889 : Quindumbo, Benguela dist. 

Specimens 

3 (J 1 $ 3 imm. 1 juv. $ Mt. Moco 6,000-7,000 feet, i5-24th Aug. (and one in 
spirit) . 

Field Notes 

This Twin-spot was found to be common on Mt. Moco at all levels, in the long 
grass of the steep mountain gullies along with Cisticola emini. 



283. (a) Estrilda astrild niediecki Reichenow, Orn. Mon. 24, p. 168, 1916 : 
Namwala, NW. Rhodesia. 

(b) E. astrild angolensis Reichenow 1902 : Malanje. 

Specimens 

(a) 1 (J Vouga, nth Aug. (b) 1 <$ 1 Mt. Moco 6,000 feet. Wing (a) 1 <J 50. (b) 
1 (J 49 : tail (a) 1 (J 51, (b) 1 $ c. 42 (moult) 41 mm. 

Systematic Notes 

The Vouga Ruddy Waxbill has light barring and a long tail and is readily identified 
with the race niediecki (synonym ngamiensis Roberts) of Barotseland and Bechuana- 
land rather than with the short-tailed, heavily barred angolensis. This represents 
a considerable extension to the known range of the southern race. 



45Q THE ECOLOGY AND TAXONOMY OF SOME ANGOLA BIRDS 

299. Fringillaria capensis nebularum Rudebeck, Bull. B.O.C. 78, p. 129, 1958 : 
Lucira, 130 m. N. of Mocamedes. 

Specimen 

1 $ Leba, 1st Sept. 

Systematic Notes 

This specimen was found to match well with the type series of the new Angola 
race. 



BIBLIOGRAPHY 

Benson, C. W. 1956. The relationship of Passer griseus (Vieillot) and P. diffusus (Smith). 
Bull. B.O.C. 76 : 38-42. 

1957- Check List of the Birds of Northern Rhodesia. 

Bocage J. V. du B. 1 88 1. Ornithologie d' Angola. 

Bo wen, W. W. 1930. Relationships and distributions of the Bare-throated Francolins 

(Pternistis). Proc. Acad. Nat. Sci. Philad. 82 : 149-164. 
Braun, R. H. 1956. Beitrag zur Biologie von Xenocopsychus ansorgei Hartert. Journ. f. 

Orn. 97 : 41-43. 
Chapin, J. P. 1932. Birds of the Belgian Congo, Pt. I. 

1939. Ibid., Pt. II. 

1953. Ibid., Part III. 

1954. Ibid., Part IV. 

1952. Campethera cailliautii and permista are conspecific. Ibis, 94 : 535. 

1948. Variation and hybridization among the Paradise Flycatchers of Africa. Evolution, 

2 : 111-126. 

Clancey, P. A. 1956. The South African races of the Black-collared Barbet Lybius torquatus 
(Dumont). Durb. Mus. Nov. 4: 273-280. 

1958. On Barbatula bocagei Sousa, 1886 : Caconda, Angola. Bull. B. O. C. 78 : 62. 

Gossweiler, J. V. & Mendonca, F. A. 1939. Carta Fitogeografica de Angola. 

Grant, C. H. B. & Mackworth-Praed, C. W. 1957. On Tchitrea melampyra Verreaux, in 
Hartlaub. Orn. Westafr., p. 90, 1857, and T. rufocinerea (Cabanis) /. /. O., p. 236, 1875. 
Bull. B. O. C. 77 : 64. 

1958. On Malaconotus blanchoti Stephens. Ibid. 78 : 19. 

Hall, B. P. 1954. A review of the Boubou Shrike, Laniarius ferrugineus . Ibis, 96 : 343-355- 

1956. Notes on a small collection of birds from Panda Matenga, N. E. Bechuanaland. 

Ostrich, 27 : 96-109. 

1958. A new race of honeyguide from Mount Moco. Bull. B. O. C. 78 : 151-152. 

1958. Variation in the Angola Lark, Mirafra angolensis Bocage. Ibid. 78 : 152-154. 

1958. A new race of Cossypha natalensis. Ibid. 78 : 154-155. 

1959- The Plain-backed Pipits of Angola. Ibid. 79 : 113-116. 

i960. Variation in the African Black Tits, Parus niger and Parus leucomelas. Ibis, 

102 : 116-123. 

& Traylor, M. A. 1959. The systematics of the African Grey Tits, Parus afer and 

P. griseiventris. Bull. B. O. C. 79 : 42-46. 

Heinrich, G. 1958. Zur Verbreitung und Lebensweise der Vogel von Angola. Journ. f. 

Orn. 99 : 121-141, 322-362, 399-421. 
Hoesch, W. & N